Neanderthal Populations Research Articles

Correlation between dental microwear analysis and dietary habits of Neanderthal populations in the Iberian Peninsula

Abstract The dietary habits of Neanderthals are considered an issue of great interest in the literature and have opened an important number of fruitful debates. Indeed, understanding diets can provide important information regarding issues of palaeoenvironmental reconstructions and subsistence strategies. In this respect, dental remains can play a vital role in the conducted efforts to reconstruct the palaeoecological niches securely and accurately since dental microwear analyses have precisely detected dietary patterns of the populations in the past. In this context, the Iberian Peninsula forms an interesting model for examining Neanderthal populations, their subsistence strategies, and adaptive skills. This study aims the examination of already published data in order to provide a holistic approach regarding the dietary habits of H. neanderthalensis populations in the Iberian Peninsula, along with the importance of the utilization of dental microwear analysis in the archaeological record.

Genetic Evidence for Geographic Structure within the Neanderthal Population

Genetic Evidence for Geographic Structure within the Neanderthal Population

Recurrent gene flow between Neanderthals and modern humans over the past 200,000 years.

Although it is well known that the ancestors of modern humans and Neanderthals admixed, the effects of gene flow on the Neanderthal genome are not well understood. We develop methods to estimate the amount of human-introgressed sequences in Neanderthals and apply it to whole-genome sequence data from 2000 modern humans and three Neanderthals. We estimate that Neanderthals have 2.5 to 3.7% human ancestry, and we leverage human-introgressed sequences in Neanderthals to revise estimates of Neanderthal ancestry in modern humans, show that Neanderthal population sizes were significantly smaller than previously estimated, and identify two distinct waves of modern human gene flow into Neanderthals. Our data provide insights into the genetic legacy of recurrent gene flow between modern humans and Neanderthals.

The Neanderthal niche space of Western Eurasia 145 ka to 30 ka ago

Neanderthals occupied Western Eurasia between 350 ka and 40 ka ago, during the climatically volatile Pleistocene. A key issue is to what extent Neanderthal populations expanded into areas of Western Eurasia and what conditions facilitated such range expansions. The range extent of Neanderthals is generally based on the distribution of Neanderthal material, but the land-altering nature of glacial periods has erased much of the already sparse material evidence of Neanderthals, particularly in the northern latitudes. To overcome this obstacle species distribution models can estimate past distributions of Neanderthals, however, most implementations are generally constrained spatially and temporally and may be artificially truncating the Neanderthal niche space. Using dated contexts from Neanderthal sites from across Western Eurasia, millennial-scale paleoclimate reconstructions, and a spatiotemporal species distribution model, we infer the fundamental climatic niche space of Neanderthals and estimate the extent of Neanderthal occupation. We find that (a.) despite the long timeframe, Neanderthals occupy a relatively narrow fundamental climatic niche space, (b.) the estimated projected potential Neanderthal niche space suggests a larger geographic range than the material record suggests, and (c.) that there was a general decline in the size of the projected potential Neanderthal niche from 145 ka ago onward, possibly contributing to their extinction.

New data on Neanderthal behavior in the Altai region, Russia

Until recently, only two sites yielding Micoquian/Keilmessergruppen (KGM) lithic assemblages were known in the Russian Altai Region, which are the result of Late European Neanderthal migrations from Eastern Europe. European Micoquian/KMG sites, often located in close proximity to one another, vary functionally, reflecting complex behavioral patterns of Neanderthal populations. Conversely, two sites in the Altai Region are identified as base camps only, suggesting that sites with other functions either have yet been undiscovered or destroyed by post-depositional processes. Here, we present new data from Verkhnyaya Sibiryachikha Cave, located close to Okladnikov Cave in southern Siberia. A stone tool typical of the Micoquian/KMG was recovered from Verkhnyaya Sibiryachikha Layers 3, chronologically overlapping Neanderthal cultural strata uncovered in Okladnikov Cave. The tool's typology and chronology suggest that it belonged to a Neanderthal from Okladnikov Cave. Verkhnyaya Sibiryachikha Cave may have been used as a game observation point for hunters pursuing prey in the river valley below. Like their European cousins, late Neanderthals in the Altai Region explored areas near their habitations, which suggests no significant changes in the behavior of the Neanderthal population that migrated eastward into Siberia.

Human evolution: Neanderthal footprints in African genomes

Human and Neanderthal populations met and mixed on multiple occasions over evolutionary time, resulting in the exchange of genetic material. New genomic analyses of diverse African populations reveal a history of bidirectional gene flow and selection acting on introgressed alleles.

Giant limpets in southern Iberian coastal and continental archaeological sites, from Neanderthals to Copper Age

The use of coastal resources has been crucial for human diet and social behaviour evolution has been extensively documented since the Middle Palaeolithic, mainly in the western Mediterranean and southern Africa. In southern Iberia, the mollusc assemblages associated with archaeological sites show a continuous record regardless of palaeoclimatic conditions. Among these, limpets are uninterruptedly abundant from the Middle Palaeolithic to the Bronze age with the giant limpets (Cymbula safiana and Patella ferruginea) being present since MIS6. To assess their distribution, predominance, and cultural significance this paper presents the results from an exhaustive archeozoological survey of the southern Iberian region. A total of 1628 specimens have been analysed, including 425 modern (333 C. safiana and 91 P. ferruginea) and 1203 from 60 archaeological deposits deriving from 36 sites, both coastal and continental. Giant limpets’ context also has been described and it has been related with a wide variety of deposits ranging from food refuse accumulations to occupations and burials.Detailed morphological and surface analysis allowed us to determine extraction methods and anthropic transformations. On coastal sites, the giant limpet record starts with Neanderthal populations from the Middle Palaeolithic and continues throughout the Upper Palaeolithic, Mesolithic, Neolithic, and Copper Age. On inland sites, giant limpets appear first in reduced numbers during the Late Neolithic and have been often worked into elliptical contours with polished, rib-free, external shell surfaces. In this study, we focus on the Copper Age, when giant limpet use becomes common, and records include tholoi, dolmens and other funerary contexts. The data indicate that the use of P. ferruginea is associated with Los Millares and El Argar cultures. Giant limpets use declines during the Bronze Age probably due to the major cultural shift that occurred during the Chalcolithic to Bronze transition.The data suggest that giant limpets should be considered as proxies of long-range trade during the Chalcolithic. We propose a new hypothesis linking giant limpets symbology with orbital/tidal variations and the ubiquitous use of marine elements as symbolism during this period.Our study highlights the relevance of shells in the reconstruction of the symbolic evolution of prehistoric cultures, as well as their connections with local ecosystem resources and regional commerce networks.

Herbivore enamel carbon and oxygen isotopes demonstrate both Homo sapiens and Neanderthals exploited similar habitats in the Zagros Mountains

ABSTRACTThe extinction of Neanderthal populations has been attributed to the onset of cold and dry climatic conditions during Marine Isotope Stage 3 or their competition with anatomically modern humans for large game resources. However, decoupling climate from competition has long proved difficult. Loess sequences and pollen cores provide regional‐scale environmental information but are less well‐suited to providing local‐scale habitat information contemporaneous with hominin habitation of occupation sites. The relationship between climate and resource availability is particularly unknown in the Zagros mountain range where archaeological evidence for both Neanderthals and Homo sapiens occupation is documented. Here, we analyse carbon (δ13C) and oxygen (δ18O) stable isotopes measured from herbivore tooth enamel carbonates recovered from the Neanderthal and modern human occupation sites of Bawa Yawan Rockshelter and Shanidar Cave to trace local‐scale floral biome dynamics and climate conditions that influence the distribution and availability of large prey targeted by both hominin species. Shared isotopic composition of herbivorous fauna, largely represented by wild goats, from both sites spanning Neanderthal and Homo sapiens occupation indicate both hominin species exploited similar habitats during climatically similar phases.

Late Pleistocene Neanderthal exploitation of stable and mosaic ecosystems in northern Iberia shown by multi-isotope evidence

AbstractDuring the last glacial period, rapidly changing environments posed substantial challenges to Neanderthal populations in Europe. Southern continental regions, such as Iberia, have been proposed as important climatic “buffer” zones during glacial phases. Contextualising the climatic and ecological conditions Neanderthals faced is relevant to interpreting their resilience. However, records of the environments and ecosystems they exploited across Iberia exhibit temporal and spatial gaps in coverage. Here we provide new evidence for palaeotemperatures, vegetation structure, and prey herbivore ecology during the late Pleistocene (MIS 5–3) in northern Spain, by applying multiple stable isotope tracers (δ18O, δ13C, δ15N, δ34S) to herbivore skeletal remains associated with Neanderthal occupations at Axlor Cave, Bizkaia. The results show little change over time and indicate stable climatic conditions and ecosystems across different occupations. Large within-layer isotopic variability in nitrogen and sulphur suggests the presence of a mosaic environment and a variety of isotopic ecotones that were exploited by Neanderthals and their prey. We implement a combination of carbonate and phosphate δ18O measurements to estimate palaeotemperatures using a cost-effective workflow. We show that the targeted use of phosphate δ18O measurements to anchor summer peak and winter trough areas enables high-precision seasonal palaeoclimatic reconstructions.

Minds on Fire: Cognitive Aspects of Early Firemaking and the Possible Inventors of Firemaking Kits

Thus far, most researchers have focused on the cognition of fire use, but few have explored the cognition of firemaking. With this contribution we analyse aspects of the two main hunter-gatherer firemaking techniques—the strike-a-light and the manual fire-drill—in terms of causal, social and prospective reasoning. Based on geographic distribution, archaeological and ethnographic information, as well as our cognitive interpretation of strike-a-light firemaking, we suggest that this technique may well have been invented by Neanderthal populations in Eurasia. Fire-drills, on the other hand, represent a rudimentary form of a symbiotic technology, which requires more elaborate prospective and causal reasoning skills. This firemaking technology may have been invented by differentHomo sapiensgroups roaming the African savanna before populating the rest of the globe, where fire-drills remain the most-used hunter-gatherer firemaking technique.

Ecological evaluation of the development of Neanderthal niche exploitation

Understanding the ecological niche occupied by Neanderthals and their ancestors is at the forefront of many Palaeolithic investigations. In this study, a complex characterization of the environments once occupied by these hominins in Western Eurasia were performed. It included the determination of the biome and climatic zones occupied, the potential wind exposures, the classification of the karst regions where they lived, the direction of the entrance of caves inhabited and the average daily total solar irradiation values. In addition, the climatic suitability of the Mediterranean mosquito fauna, tick-borne encephalitis and four large mammal species as potential mammal hosts were also studied. It was found that Western Eurasian hominins occupied a wide ecological niche range from the semi-arid to the subarctic climates. Between the MIS20-MIS4 interval, the biomes occupied by Neanderthals and their ancestors shifted from the woodland-shrubland to the boreal biomes, indicating the increasing general biocultural adaptation to the more continental environments. These populations inhabited territories with weak to moderate wind exposure (Beaufort 2–5) and usually occupied caves with a south or southwest-oriented entrance. This habitat selection indicates the recognition of local microclimatic factors and a conscious choice of shelters. In the glacial periods, the increasing extension of temperate-subpolar karst regions at the expense of the more habitable subtropical karst areas might have adversely affected the overwintering chances of ancient western Eurasian populations in West, Central Europe, and Central Asia. The supraorbital area of ancient hominins with low receding foreheads and prominent supraorbital ridges may have been exposed to a greater risk of the development of frontal sinusitis and its complications than modern humans with more gracile skulls. Neanderthals and their predecessors were exposed to moderate average daily solar radiation values, although these values exhibit heterogenic picture in the comparison of Central and Southern Europe. While the southern European Neanderthal populations were dominantly exposed to the risk of mosquito-borne diseases, the northern and central populations could suffer from tick-borne diseases. However, both tick-borne encephalitis virus and its medium-large mammal hosts were present in most sites dwelled by Neanderthals and their ancestors.

Assessing climatic impact on transition from Neanderthal to anatomically modern human population on Iberian Peninsula: a macroscopic perspective

The Iberian Peninsula is of particular interest for the research on the Neanderthal (NEA) to anatomically modern human (AMH) population transition. The AMHs arrived in Iberia last from Eastern Europe and thus any possible contacts between the two populations occurred here later than elsewhere. The transition process took place in the earlier part of the Marine Isotope Stage 3 (∼60–27 cal ka BP) as repeated and profound climate changes challenged the population stability. To investigate how climate change and population interactions influenced the transition, we combine climate data with archaeological-site data to reconstruct the Human Existence Potential, a measure of the probability of human existence, for both the NEA and AMH populations in the Greenland Interstadial 11–10 (GI11-10) and Stadial 10–9/Heinrich event 4 (GS10-9/HE4) times. It is found that during GS10-9/HE4, large parts of the peninsula became unsuitable for NEA human existence and the NEA settlement areas contracted to isolated coastal hot spots. As a consequence, the NEA networks became highly unstable, triggering the final collapse of the population. The AMHs arrived in Iberia in GI10 but were confined to patches in the northern most strip of the peninsula. They were soon facing the much colder climate of GS10-9/HE4, which prevented their further expansion or even caused a contraction of their settlement areas. Thus, due to the constellation of climate change and the dispersal of the two populations into different regions of the peninsula, it is unlikely that the NEAs and AMHs coexisted in extensive areas and the AMHs had a significant influence on the demography of the NEAs.

Archaeological evidence for two culture diverse Neanderthal populations in the North Caucasus and contacts between them.

Neanderthals were widespread during the Middle Palaeolithic (MP) across Europe and Asia, including the Caucasus Mountains. Occupying the border between eastern Europe and West Asia, the Caucasus is important region regarding the Neanderthal occupation of Eurasia. On current radiometric estimates, the MP is represented in the Caucasus between about 260-210 ka and about 40 ka. Archaeological record indicates that several culture diverse MP hominin populations inhabited the Caucasus, but the region complex population history during this period remains poorly understood. In this paper, we identify for the first time the archaeological evidence indicating contacts between two culture diverse MP Neanderthal populations in the North Caucasus and discuss the nature of these contacts. Basing on the lithic assemblages that we excavated at Mezmaiskaya cave in the north-western Caucasus (Kuban River basin) and Saradj-Chuko grotto in the north-central Caucasus (Terek River basin), dating from MIS 5 to MIS 3, and comparative data from other MP sites in the Caucasus, we identify two large cultural regions that existed during the late MP in the North Caucasus. The distinctive toolkits and stone knapping technologies indicate that the MP assemblages from Mezmaiskaya cave and other sites in the west of North Caucasus represent a Caucasian variant of the Eastern Micoquian industry that was wide spread in central and eastern Europe, while the assemblages from Saradj-Chuko Grotto and other sites in the east of North Caucasus closely resemble the Zagros Mousterian industry that was wide spread in the Armenian Highlands, Lesser Caucasus and Zagros Mountains. The archaeological evidence implies that two culture diverse populations of Neanderthals settled the North Caucasus during the Late Pleistocene from two various source regions: from the Armenian Highlands and Lesser Caucasus along the Caspian Sea coast, and from Russian plain along the Sea of Azov coast.

A multi-proxy study from new excavations in the Middle Palaeolithic site of Cova del Puntal del Gat (Benirredrà, València, Spain)

The Mediterranean basin constitutes one of the best areas to analyse Neanderthal populations and lifestyles in South-Western Europe. In this context, new excavations conducted in the Middle Palaeolithic site of Cova del Puntal del Gat expand the information available regarding this rich region. In this study, new results are reported, including detailed studies on stratigraphy, lithic technology, anthracology, carpology, and zooarchaeology and taphonomy of macro and micromammals, with the final objective of characterizing the Neanderthals’ subsistence strategies and occupational patterns. These results are framed within a broader regional study perspective that includes MIS 5 and 4 sites. Chronostratigraphic review has enabled us to reorganize many sites that were originally included in MIS 3, towards older stages belonging to the end of MIS 4 and throughout MIS 5.

Virtual excavation and analysis of the early Neanderthal cranium from Altamura (Italy)

Complete Neanderthal skeletons are almost unique findings. A very well-preserved specimen of this kind was discovered in 1993 in the deepest recesses of a karstic system near the town of Altamura in Southern Italy. We present here a detailed description of the cranium, after we virtually extracted it from the surrounding stalagmites and stalactites. The morphology of the Altamura cranium fits within the Neanderthal variability, though it retains features occurring in more archaic European samples. Some of these features were never observed in Homo neanderthalensis, i.e. in fossil specimens dated between 300 and 40 ka. Considering the U-Th age we previously obtained (>130 ka), the morphology of Altamura suggests that the archaic traits it retains may have been originated by geographic isolation of the early Neanderthal populations from Southern Italy.

Neanderthal subsistence at Chez-Pinaud Jonzac (Charente-Maritime, France): A kill site dominated by reindeer remains, but with a horse-laden diet?

During the MIS 4 in Southwestern France, Quina Neanderthal from the north of the Aquitaine was characterized by a hunting specialization on the reindeer and the lack of diversity in their diet. They developed task-specific locations dedicated to the capture, the butchery, and the consumption of reindeer, and the whole society seems, in this region, to be dependent on this food resource. In this context, the site of Chez-Pinaud at Jonzac (France) occupies a specific place. First, interpreted as a reindeer kill and butchery site, the recent recovery of the site underlines the importance of the large ungulate (horse and bison) to the faunal spectrum (30% of the NISP). Considering the quantity of meat and grease that these species can provide to hunters, the new zooarchaeological analyses suggest that at least the horse may have played a major role in the diet of the Neanderthal population. Since Jonzac is one of the largest sites for this period, these results relativize the importance of reindeer specialization of the Quina population and the lack of diversityl in their diet.

A long-term perspective on Neanderthal environment and subsistence: Insights from the dental microwear texture analysis of hunted ungulates at Combe-Grenal (Dordogne, France).

Large bovids and cervids constituted major components of the European Middle Palaeolithic faunas and hence a key resource for Neanderthal populations. In paleoenvironmental reconstructions, red deer (Cervus elaphus) occurrence is classically considered as a tree-cover indicator while Bovinae (Bison priscus and Bos primigenius) and reindeer (Rangifer tarandus) occurrences are typically associated with open landscapes. However, insights into the ecology of extant ungulate populations show a more complex reality. Exploring the diet of past ungulates allows to better comprehend the hunting strategies of Palaeolithic populations and to reconstruct the modifications through time of past landscapes. By reflecting what animals have eaten during the last days or weeks of their life, dental microwear textures of herbivores link a population and its environment. Here we analyzed, via Dental Microwear Texture Analysis (DMTA), the diet of 50 Bos/Bison, 202 R. tarandus and 116 C. elaphus preyed upon by the Neanderthals that occupied Combe-Grenal rock-shelter, one of the most important Mousterian archaeo-sequences in southwestern France considering its long stratigraphy, abundance of faunal remains and the variations perceptible in Palaeolithic material culture. Grazers and mixed-feeders are the most represented dietary categories among Combe-Grenal's guild of herbivores, highlighting the availability, along the sequence, of open landscapes. The absence of clear changes in the use of plant resources by hunted ungulates through time, even though palaeoenvironmental changes were well-documented by previous studies along the sequence, is interpreted as resulting from the hunting of non-randomly selected prey by Neanderthals, preferentially in open environments. Thus, these results provide further insight into the hunting strategies of Neanderthals and modify our perception of potential links between subsistence and material culture. Combe-Grenal hunters "stayed in the open" through millennia, and were not forced to switch to hunting tactics and material technology adapted to close encounters in forested environments.

A pedogenically-informed chronostratigraphic model elucidates the geochronology at the type site of the Bohunician technocomplex

The Bohunician, a Moravian variant of the broader Initial Upper Palaeolithic (IUP) lithic technocomplex, is thought to have arisen during the transition from the Middle to the Upper Palaeolithic in the ‘contact zone’ between the first anatomically modern humans and local Neanderthals. Attempts to date the Bohunician occupation at the eponymous site of Brno-Bohunice using luminescence and radiocarbon methods have yielded contradictory results. We propose a new perspective of the chronological position and taphonomy of the Bohunician finds based on a revaluation of published radiometric and luminescence data from the Brno-Bohunice site according to an updated radiocarbon calibration curve and recently revised pedostratigraphy. Here, for the first time, we combined contradictionary results of various dating method into one chronostratigraphical model of the site. The Bayesian model considers the discovery that the palaeosol(s) contemporary to most of the artefacts is (are) missing and the actual position was affected by frost-heaving processes. The onset of the Bohunician technocomplex is therefore placed most probably as early as ∼50 ka cal BP (the end of GI 14) and shows that it existed mainly during the interstadials GI 13 and GI 12. Nevertheless, the broad timespan of unmodelled thermoluminescence dates indicates a possible persistence of the last practitioners of Bohunician technology until ∼42 ka cal BP (GI 11), when development of a Cambisol horizon took place. Our model has two possible explanations: The first wave of anatomically modern humans (with or without Neanderthal introgression) arrived in Central Europe before GI 12, or the Bohunician technocomplex is connected with the local Neanderthal population.

Genetic insights into the social organization of Neanderthals

Genomic analyses of Neanderthals have previously provided insights into their population history and relationship to modern humans1–8, but the social organization of Neanderthal communities remains poorly understood. Here we present genetic data for 13 Neanderthals from two Middle Palaeolithic sites in the Altai Mountains of southern Siberia: 11 from Chagyrskaya Cave9,10 and 2 from Okladnikov Cave11—making this one of the largest genetic studies of a Neanderthal population to date. We used hybridization capture to obtain genome-wide nuclear data, as well as mitochondrial and Y-chromosome sequences. Some Chagyrskaya individuals were closely related, including a father–daughter pair and a pair of second-degree relatives, indicating that at least some of the individuals lived at the same time. Up to one-third of these individuals’ genomes had long segments of homozygosity, suggesting that the Chagyrskaya Neanderthals were part of a small community. In addition, the Y-chromosome diversity is an order of magnitude lower than the mitochondrial diversity, a pattern that we found is best explained by female migration between communities. Thus, the genetic data presented here provide a detailed documentation of the social organization of an isolated Neanderthal community at the easternmost extent of their known range.

A Case of Chronic Maxillary Sinusitis in a Late Neanderthal Population of the Altai Mountains

We describe a likely case of chronic maxillary sinusitis (CMS) in a Neanderthal skeletal sample from Chagyrskaya Cave, in the Altai Mountains. Signs of CMS were recorded in the Chagyrskaya 57 specimen, which is a fragment of a left maxilla. Alveoli of the upper fi rst molar are partially preserved, and so are the second and third upper molars, with adjacent parts of the walls, and the fl oor of the maxillary sinus. The fragment was found in layer 6b, dating to 53,100–51,100 BP. We analyze the factors that had caused the development of the disease, and assess its etiology. In the 3D-model, generated by computed microtomography, and in the original specimen, porotic changes were registered, situated at the fracture line of the alveoli of M1, lost post-mortem, and near the vestibular roots of both preserved molars. Also, there were isolated bone spicules, 1.0–2.6 mm in size. These signs indicate incipient CMS, evidently caused by chronic periodontal disease combined with a deep alveolar recess of the maxillary sinus. As the periodontal gap expanded, several small nutrient foramina, piercing the bottom of the sinus, merged. As a result, several oro-antral channels formed, whereupon the infection spread into the maxillary sinus. Since the deep alveolar recess is observed in the vast majority of Neanderthal crania with published images or reconstructed maxillary cavities, it can be assumed that Neanderthals were predisposed to odontogenic CMS.