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Abstract The emergence of the Middle Palaeolithic, and its variability over time and space are key questions in the field of prehistoric archaeology. Many sites have been documented in the south-eastern margins of the Massif central and the middle Rhône valley, a migration path that connects Northern Europe with the Mediterranean. Well-dated, long stratigraphic sequences are essential to understand Neanderthals dynamics and demise, and potential interactions with Homo sapiens in the area, such as the one displayed at the Maras rock shelter (“Abri du Maras”). The site is characterised by exceptional preservation of archaeological remains, including bones dated using radiocarbon (14C) and teeth using electron spin resonance combined with uranium series (ESR/U-series). Optically stimulated luminescence was used to date the sedimentary deposits. By combining the new ages with previous ones using Bayesian modelling, we are able to clarify the occupation time over a period spanning 200,000 years. Between ca. 250 and 40 ka, the site has been used as a long-term residence by Neanderthals, specifically during three interglacial periods: first during marine isotopic stage (MIS) 7, between 247 ± 34 and 223 ± 33 ka, and then recurrently during MIS 5 (between 127 ± 17 and 90 ± 9 ka) and MIS 3 (up to 39,280 cal BP).

Abstract The phylogenetic position of Homo habilis is central to debates over the origin and early evolution of the genus Homo. A large portion of the species hypodigm consists of dental remains, but they have only been studied at the often worn enamel surface. We investigate the morphology of the H. habilis enamel-dentine junction (EDJ), which is preserved in cases of moderate tooth wear and known to carry a strong taxonomic signal. Geometric morphometrics is used to characterise dentine crown shape and size across the entire mandibular and maxillary tooth rows, compared with a broad comparative sample (n = 712). We find that EDJ morphology in H. habilis is for the most part remarkably primitive, supporting the hypothesis that the H. habilis hypodigm has more in common with Australopithecus than later Homo. Additionally, the chronologically younger specimen OH 16 displays a suite of derived features; its inclusion in H. habilis leads to excessive levels of variation.

Abstract High-throughput proteomic analysis of archaeological skeletal remains provides information about past fauna community compositions and species dispersals in time and space. Archaeological skeletal remains are a finite resource, however, and therefore it becomes relevant to optimize methods of skeletal proteome extraction. Ancient proteins in bone specimens can be highly degraded and consequently, extraction methods for well-preserved or modern bone might be unsuitable for the processing of highly degraded skeletal proteomes. In this study, we compared six proteomic extraction methods on Late Pleistocene remains with variable levels of proteome preservation. We tested the accuracy of species identification, protein sequence coverage, deamidation, and the number of post-translational modifications per method. We find striking differences in obtained proteome complexity and sequence coverage, highlighting that simple acid-insoluble proteome extraction methods perform better in highly degraded contexts. For well-preserved specimens, the approach using EDTA demineralization and protease-mix proteolysis yielded a higher number of identified peptides. The protocols presented here allowed protein extraction from ancient bone with a minimum number of working steps and equipment and yielded protein extracts within three working days. We expect further development along this route to benefit large-scale screening applications of relevance to archaeological and human evolution research.

Abstract The evolution of the medial longitudinal arch (MLA) is one of the most impactful adaptations in the hominin foot that emerged with bipedalism. When and how it evolved in the human lineage is still unresolved. Complicating the issue, clinical definitions of flatfoot in living Homo sapiens have not reached a consensus. Here we digitally investigate the navicular morphology of H. sapiens (living, archaeological, and fossil), great apes, and fossil hominins and its correlation with the MLA. A distinctive navicular shape characterises living H. sapiens with adult acquired flexible flatfoot, while the congenital flexible flatfoot exhibits a ‘normal’ navicular shape. All H. sapiens groups differentiate from great apes independently from variations in the MLA, likely because of bipedalism. Most australopith, H. naledi, and H. floresiensis navicular shapes are closer to those of great apes, which is inconsistent with a human-like MLA and instead might suggest a certain degree of arboreality. Navicular shape of OH 8 and fossil H. sapiens falls within the normal living H. sapiens spectrum of variation of the MLA (including congenital flexible flatfoot and individuals with a well-developed MLA). At the same time, H. neanderthalensis seem to be characterised by a different expression of the MLA.

Abstract Secure environmental contexts are crucial for hominin interpretation and comparison. The discovery of a Denisovan individual and associated fauna at Tam Ngu Hao 2 (Cobra) Cave, Laos, dating back to 164–131 ka, allows for environmental comparisons between this (sub)tropical site and the Palearctic Denisovan sites of Denisova Cave (Russia) and Baishiya Karst Cave (China). Denisovans from northern latitudes foraged in a mix of forested and open landscapes, including tundra and steppe. Using stable isotope values from the Cobra Cave assemblage, we demonstrate that, despite the presence of nearby canopy forests, the Denisovan individual from Cobra Cave primarily consumed plants and/or animals from open forests and savannah. Using faunal evidence and proxy indicators of climates, results herein highlight a local expansion of rainforest at ~ 130 ka, raising questions about how Denisovans responded to this local climate change. Comparing the diet and habitat of the archaic hominin from Cobra Cave with those of early Homo sapiens from Tam Pà Ling Cave (46–43 ka), Laos, it appears that only our species was able to exploit rainforest resources.

Abstract Here we present stable carbon, nitrogen and sulfur isotope ratios of collagen extracted from Rangifer, Equus and Bison bone (n = 128) from different stratigraphic levels at the chronologically well-constrained Middle and Upper Palaeolithic site of Les Cottés, France. Samples were taken from five phases of site use (US08, US06, US04 [upper and lower], and US02; ~ 45.8–35.3 ka cal BP) to explore the dietary and spatial palaeoecology of these ungulate species during MIS 3, and the contemporary climate. Temporal trends in δ 15N values of all species broadly align with other climatic indicators at the site and the lowest values in US04 correspond to the Heinrich 4 cooling event, reflecting changes in the composition of soil/plant nitrogen at this time. Rangifer collagen is 13C-enriched compared to the other species throughout, consistent with lichen consumption. However, this isotopic niche partitioning between Rangifer and Equus/Bison is most extensive during US04, indicating plasticity in reindeer feeding behaviour, and potentially overall increased lichen biomass during this cooler/more arid phase. Rangifer δ 34S values are consistently lower than Equus and Bison, which could be indicative of their more extensive spatial ranges incorporating greater inland areas. Equus and Bison demonstrate a significant decrease in δ 34S values through time, which may be linked to contemporary climatic decline.

Abstract Around 42,000 years ago, anatomically modern humans appeared in Western Europe to the detriment of indigenous Neanderthal groups. It is during this period that new techno-cultural complexes appear, such as the Châtelperronian that extends from northern Spain to the Paris Basin. The Grotte du Renne (Arcy-sur-Cure) is a key site for discussing the biological identity of its makers. This deposit has yielded several Neanderthal human remains in its Châtelperronian levels. However, the last inventory of the paleoanthropological collection attributed to this techno-complex allowed the identification of an ilium belonging to a neonate (AR-63) whose morphology required a thorough analysis to assess its taxonomic attribution. Using geometric morphometrics, we quantified its morphology and compared it to that of 2 Neanderthals and 32 recent individuals deceased during the perinatal period to explore their morphological variation. Our results indicate a morphological distinction between the ilia of Neanderthals and anatomically modern neonates. Although AR-63 is slightly outside recent variability, it clearly differs from the Neanderthals. We propose that this is due to its belonging to an early modern human lineage whose morphology differs slightly from present-day humans. We also explore different hypotheses about the presence of this anatomically modern neonate ilium among Neanderthal remains.

[This corrects the article DOI: 10.1371/journal.pone.0254848.].

The dispersal of Homo sapiens across Eurasia during MIS 3 in the Late Pleistocene is marked by technological shifts and other behavioral changes, known in the archaeological record under the term of Initial Upper Paleolithic (IUP). Bacho Kiro Cave in north Bulgaria, re-excavated by us from 2015 to 2021, is one of the reference sites for this phenomenon. The newly excavated lithic assemblages dated by radiocarbon between 45,040 and 43,280 cal BP and attributed to Homo sapiens encompass more than two thousand lithic artifacts. The lithics, primarily from Layer N1-I, exist amid diverse fauna remains, human fossils, pierced animal teeth pendants, and sediment with high organic content. This article focuses on the technological aspects of the IUP lithics, covering raw material origin and use-life, blank production, on-site knapping activities, re-flaking of lithic implements, and the state of retouched lithic components. We apply petrography for the identification of silicites and other used stones. We employ chaîne opératoire and reduction sequence approaches to profile the lithics techno-typologically and explore the lithic economy, particularly blade production methods, knapping techniques, and artifact curation. Raw material analysis reveals Lower Cretaceous flints from Ludogorie and Upper Cretaceous flints from the Danube region, up to 190 km and 130 km, respectively, from Bacho Kiro Cave, indicating long-distance mobility and finished products transport. Imported lithic implements, were a result of unidirectional and bidirectional non-Levallois laminar technology, likely of volumetric concept. Systematic on-anvil techniques (bipolar knapping) and tool segmentation indicate re-flaking and reshaping of lithic implements, reflecting on-site curation and multifaceted lithic economy. A limited comparison with other IUP sites reveals certain shared features and also regional variations. Bacho Kiro Cave significantly contributes to understanding the technological and behavioral evolution of early Homo sapiens in western Eurasia.

Abstract Figurative depictions in art first occur ca. 50,000 years ago in Europe, Africa, and Southeast Asia. Considered by most as an advanced form of symbolic behavior, they are restricted to our species. Here, we report a piece of ornament interpreted as a phallus-like representation. It was found in a 42,000 ca.-year-old Upper Paleolithic archaeological layer at the open-air archaeological site of Tolbor-21, in Mongolia. Mineralogical, microscopic, and rugosimetric analyses points toward the allochthonous origin of the pendant and a complex functional history. Three-dimensional phallic pendants are unknown in the Paleolithic record, and this discovery predates the earliest known sexed anthropomorphic representation. It attests that hunter-gatherer communities used sex anatomical attributes as symbols at a very early stage of their dispersal in the region. The pendant was produced during a period that overlaps with age estimates for early introgression events between Homo sapiens and Denisovans, and in a region where such encounters are plausible.

Abstract The timing of the first arrival of Homo sapiens in East Asia from Africa and the degree to which they interbred with or replaced local archaic populations is controversial. Previous discoveries from Tam Pà Ling cave (Laos) identified H. sapiens in Southeast Asia by at least 46 kyr. We report on a recently discovered frontal bone (TPL 6) and tibial fragment (TPL 7) found in the deepest layers of TPL. Bayesian modeling of luminescence dating of sediments and U-series and combined U-series-ESR dating of mammalian teeth reveals a depositional sequence spanning ~86 kyr. TPL 6 confirms the presence of H. sapiens by 70 ± 3 kyr, and TPL 7 extends this range to 77 ± 9 kyr, supporting an early dispersal of H. sapiens into Southeast Asia. Geometric morphometric analyses of TPL 6 suggest descent from a gracile immigrant population rather than evolution from or admixture with local archaic populations.

Summary: Six infant human teeth and 112 animal tooth pendants from Borsuka Cave were identified as the oldest burial in Poland. However, uncertainties around the dating and the association of the teeth to the pendants have precluded their association with an Upper Palaeolithic archaeological industry. Using

This study demonstrates how proximal femoral adaptations to bipedal locomotion and lifestyles impact the risk of intracapsular hip fracture, independent of osteoporosis

Abstract North Africa is a key area for understanding hominin population movements and the expansion of our species. It is home to the earliest currently known Homo sapiens (Jebel Irhoud) and several late Middle Stone Age (MSA) fossils, notably Kébibat, Contrebandiers 1, Dar-es-Soltane II H5 and El Harhoura. Mostly referred to as “Aterian” they fill a gap in the North African fossil record between Jebel Irhoud and Iberomaurusians. We explore morphological continuity in this region by quantifying mandibular shape using 3D (semi)landmark geometric morphometric methods in a comparative framework of late Early and Middle Pleistocene hominins (n = 15), Neanderthals (n = 27) and H. sapiens (n = 145). We discovered a set of mixed features among late MSA fossils that is in line with an accretion of modern traits through time and an ongoing masticatory gracilization process. In Northern Africa, Aterians display similarities to Iberomaurusians and recent humans in the area as well as to the Tighenif and Thomas Quarry hominins, suggesting a greater time depth for regional continuity than previously assumed. The evidence we lay out for a long-term succession of hominins and humans emphasizes North Africa’s role as source area of the earliest H. sapiens.

Here the authors demonstrate the use of zinc isotopes (δ66Zn) to geochemically assess trophic levels in extant and extinct sharks. They show that the Neogene megatooth shark (Otodus megalodon) and the great white shark (Carcharodon carcharias) occupied a similar trophic level.

Evidence for the presence of Homo during the Middle Pleistocene is limited in continental Southeast Asia. Here, the authors report a hominin molar from Tam Ngu Hao 2 (Cobra Cave), dated to 164–131 kyr. They use morphological and paleoproteomic analysis to show that it likely belonged to a female Denisovan.

Abstract Bone surface modifications are crucial for understanding human subsistence and dietary behaviour, and can inform about the techniques employed in the production and use of bone tools. Permission to destructively sample such unique artefacts is not always granted. The recent development of non-destructive proteomic extraction techniques has provided some alternatives for the analysis of rare and culturally significant artefacts, including bone tools and personal ornaments. The Eraser Extraction Method (EEM), first developed for ZooMS analysis of parchment, has recently been applied to bone and ivory specimens. To test the potential impact of the EEM on ancient bone surfaces, we analyse six anthropogenically modified Palaeolithic bone specimens from Bacho Kiro Cave (Bulgaria) through a controlled sampling experiment using qualitative and 3D quantitative microscopy. Although the overall bone topography is generally preserved, our findings demonstrate a slight flattening of the microtopography alongside the formation of micro-striations associated with the use of the eraser for all bone specimens. Such modifications are similar to ancient use-wear traces. We therefore consider the EEM a destructive sampling approach for Palaeolithic bone surfaces. Together with low ZooMS success rates in some of the reported studies, the EEM might not be a suitable approach to taxonomically identify Pleistocene bone specimens.

Abstract Evidence of mobiliary art and body augmentation are associated with the cultural innovations introduced by Homo sapiens at the beginning of the Upper Paleolithic. Here, we report the discovery of the oldest known human-modified punctate ornament, a decorated ivory pendant from the Paleolithic layers at Stajnia Cave in Poland. We describe the features of this unique piece, as well as the stratigraphic context and the details of its chronometric dating. The Stajnia Cave plate is a personal 'jewellery' object that was created 41,500 calendar years ago (directly radiocarbon dated). It is the oldest known of its kind in Eurasia and it establishes a new starting date for a tradition directly connected to the spread of modern Homo sapiens in Europe.

Abstract Childhood is an ontogenetic stage unique to the modern human life history pattern. It enables the still dependent infants to achieve an extended rapid brain growth, slow somatic maturation, while benefitting from provisioning, transitional feeding, and protection from other group members. This tipping point in the evolution of human ontogeny likely emerged from early Homo. The GAR IVE hemi-mandible (1.8 Ma, Melka Kunture, Ethiopia) represents one of the rarely preserved early Homo infants (~ 3 years at death), recovered in a richly documented Oldowan archaeological context. Yet, based on the sole external inspection of its teeth, GAR IVE was diagnosed with a rare genetic disease–amelogenesis imperfecta (AI)–altering enamel. Since it may have impacted the child’s survival, this diagnosis deserves deeper examination. Here, we reassess and refute this diagnosis and all associated interpretations, using an unprecedented multidisciplinary approach combining an in-depth analysis of GAR IVE (synchrotron imaging) and associated fauna. Some of the traits previously considered as diagnostic of AI can be better explained by normal growth or taphonomy, which calls for caution when diagnosing pathologies on fossils. We compare GAR IVE’s dental development to other fossil hominins, and discuss the implications for the emergence of childhood in early Homo.

Abstract The capability of Pleistocene hominins to successfully adapt to different types of tropical forested environments has long been debated. In order to investigate environmental changes in Southeast Asia during a critical period for the turnover of hominin species, we analysed palaeoenvironmental proxies from five late Middle to Late Pleistocene faunas. Human teeth discoveries have been reported at Duoi U’Oi, Vietnam (70–60 ka) and Nam Lot, Laos (86–72 ka). However, the use of palaeoproteomics allowed us to discard the latter, and, to date, no human remains older than ~ 70 ka are documented in the area. Our findings indicate that tropical rainforests were highly sensitive to climatic changes over that period, with significant fluctuations of the canopy forests. Locally, large-bodied faunas were resilient to these fluctuations until the cooling period of the Marine Isotope Stage 4 (MIS 4; 74–59 ka) that transformed the overall biotope. Then, under strong selective pressures, populations with new phenotypic characteristics emerged while some other species disappeared. We argue that this climate-driven shift offered new foraging opportunities for hominins in a novel rainforest environment and was most likely a key factor in the settlement and dispersal of our species during MIS 4 in SE Asia.

McCormack et al. present the first comparison of archaeological δ66Zn from bone material compared with traditional δ15 N and δ13 C values derived from collagen samples over a large geographic area. This study shows that the inclusion of δ66Zn analysis can provide otherwise inaccessible supplementary dietary information and more robust trophic level estimations.

Abstract Exploring the role of changing climates in human evolution is currently impeded by a scarcity of climatic information at the same temporal scale as the human behaviors documented in archaeological sites. This is mainly caused by high uncertainties in the chronometric dates used to correlate long-term climatic records with archaeological deposits. One solution is to generate climatic data directly from archaeological materials representing human behavior. Here we use oxygen isotope measurements of Bos/Bison tooth enamel to reconstruct summer and winter temperatures in the Late Pleistocene when Neandertals were using the site of La Ferrassie. Our results indicate that, despite the generally cold conditions of the broader period and despite direct evidence for cold features in certain sediments at the site, Neandertals used the site predominantly when climatic conditions were mild, similar to conditions in modern day France. We suggest that due to millennial scale climate variability, the periods of human activity and their climatic characteristics may not be representative of average conditions inferred from chronological correlations with long-term climatic records. These results highlight the importance of using direct routes, such as the high-resolution archives in tooth enamel from anthropogenically accumulated faunal assemblages, to establish climatic conditions at a human scale.

Summary: Sicily is a key region for understanding the agricultural transition in the Mediterranean because of its central position. Here, we present genomic and stable isotopic data for 19 prehistoric Sicilians covering the Mesolithic to Bronze Age periods (10,700–4,100 yBP). We find that Early Mesolithic hunter-gatherers (HGs) from Sicily are a highly drifted lineage of the Early Holocene western European HGs, whereas Late Mesolithic HGs carry ∼20% ancestry related to northern and (south) eastern European HGs, indicating substantial gene flow. Early Neolithic farmers are genetically most similar to farmers from the Balkans and Greece, with only ∼7% of ancestry from local Mesolithic HGs. The genetic discontinuities during the Mesolithic and Early Neolithic match the changes in material culture and diet. Three outlying individuals dated to ∼8,000 yBP; however, suggest that hunter-gatherers interacted with incoming farmers at Grotta dell’Uzzo, resulting in a mixed economy and diet for a brief interlude at the Mesolithic-Neolithic transition.

Abstract The origin of funerary practices has important implications for the emergence of so-called modern cognitive capacities and behaviour. We provide new multidisciplinary information on the archaeological context of the La Ferrassie 8 Neandertal skeleton (grand abri of La Ferrassie, Dordogne, France), including geochronological data -14C and OSL-, ZooMS and ancient DNA data, geological and stratigraphic information from the surrounding context, complete taphonomic study of the skeleton and associated remains, spatial information from the 1968–1973 excavations, and new (2014) fieldwork data. Our results show that a pit was dug in a sterile sediment layer and the corpse of a two-year-old child was laid there. A hominin bone from this context, identified through Zooarchaeology by Mass Spectrometry (ZooMS) and associated with Neandertal based on its mitochondrial DNA, yielded a direct 14C age of 41.7–40.8 ka cal BP (95%), younger than the 14C dates of the overlying archaeopaleontological layers and the OSL age of the surrounding sediment. This age makes the bone one of the most recent directly dated Neandertals. It is consistent with the age range for the Châtelperronian in the site and in this region and represents the third association of Neandertal taxa to Initial Upper Palaeolithic lithic technocomplex in Western Europe. A detailed multidisciplinary approach, as presented here, is essential to advance understanding of Neandertal behavior, including funerary practices.

Studies of hominin dental morphology frequently consider accessory cusps on the lower molars, in particular those on the distal margin of the tooth (C6 or distal accessory cusp) and the lingual margin of the tooth (C7 or lingual accessory cusp). They are often utilized in studies of hominin systematics, where their presence or absence is assessed at the outer enamel surface (OES). However, studies of the enamel-dentine junction (EDJ) suggest these traits may be more variable in development, morphology and position than previously thought. Building on these studies, we outline a scoring procedure for the EDJ expression of these accessory cusps that considers the relationship between these accessory cusps and the surrounding primary cusps. We apply this scoring system to a sample of Plio-Pleistocene hominin mandibular molars of Paranthropus robustus, Paranthropus boisei, Australopithecus afarensis, Australopithecus africanus, Homo sp., Homo habilis and Homo erectus from Africa and Asia (n = 132). We find that there are taxon-specific patterns in accessory cusp expression at the EDJ that are consistent with previous findings at the OES. For example, P. robustus M1s and M2s very often have a distal accessory cusp but no lingual accessory cusp, while H. habilis M1s and M2s show the opposite pattern. The EDJ also reveals a number of complicating factors; some apparent accessory cusps at the enamel surface are represented at the EDJ only by shouldering on the ridges associated with the main cusps, while other accessory cusps appear to have little or no EDJ expression at all. We also discuss the presence of double and triple accessory cusps, including the presence of a double lingual accessory cusp on the distal ridge of the metaconid in the type specimen of H. habilis (OH 7–M1) that is not clear at the OES due to occlusal wear. Overall, our observations, as well as our understanding of the developmental underpinnings of cusp patterning, suggest that we should be cautious in our comparisons of accessory cusps for taxonomic interpretations.

Neanderthal extinction has been a matter of debate for many years. New discoveries, better chronologies and genomic evidence have done much to clarify some of the issues. This evidence suggests that Neanderthals became extinct around 40,000-37,000 years before present (BP), after a period of coexistence with Homo sapiens of several millennia, involving biological and cultural interactions between the two groups. However, the bulk of this evidence relates to Western Eurasia, and recent work in Central Asia and Siberia has shown that there is considerable local variation. Southwestern Asia, despite having a number of significant Neanderthal remains, has not played a major part in the debate over extinction. Here we report a Neanderthal deciduous canine from the site of Bawa Yawan in the West-Central Zagros Mountains of Iran. The tooth is associated with Zagros Mousterian lithics, and its context is preliminary dated to between ~43,600 and ~41,500 years ago.

To investigate the mobility patterns of Neanderthals and modern humans in Europe during the Middle-to-Upper Palaeolithic transition period, we applied strontium isotope analysis to Neanderthal (n = 3) and modern human (n = 2) teeth recovered from the site of Fumane Cave in the Monti Lessini region of Northern Italy. We also measured a large number of environmental samples from the region, to establish a strontium 'baseline', and also micromammals (vole teeth) from the levels associated with the hominin teeth. We found that the modern humans and Neanderthals had similar strontium isotope values, and these values match the local baseline values we obtained for the site and the surrounding region. We conclude that both groups were utilizing the local mountainous region where Fumane Cave is situated, and likely the nearby Lessini highlands and Adige plains, and therefore the strontium evidence does not show differening mobility patterns between Neanderthals and modern humans at the Fumane site.

Abstract The late Middle Palaeolithic (MP) settlement patterns in the Levant included the repeated use of caves and open landscape sites. The fossil record shows that two types of hominins occupied the region during this period—Neandertals and Homo sapiens. Until recently, diagnostic fossil remains were found only at cave sites. Because the two populations in this region left similar material cultural remains, it was impossible to attribute any open-air site to either species. In this study, we present newly discovered fossil remains from intact archaeological layers of the open-air site ‘Ein Qashish, in northern Israel. The hominin remains represent three individuals: EQH1, a nondiagnostic skull fragment; EQH2, an upper right third molar (RM3); and EQH3, lower limb bones of a young Neandertal male. EQH2 and EQH3 constitute the first diagnostic anatomical remains of Neandertals at an open-air site in the Levant. The optically stimulated luminescence ages suggest that Neandertals repeatedly visited ‘Ein Qashish between 70 and 60 ka. The discovery of Neandertals at open-air sites during the late MP reinforces the view that Neandertals were a resilient population in the Levant shortly before Upper Palaeolithic Homo sapiens populated the region.

Peptide mass fingerprinting of bone collagen (ZooMS) has previously been proposed as a method to calculate the extent of the non-enzymatic degradation of glutamine into glutamic acid (deamidation). Temporal and spatial variation of glutamine deamidation at a single site, however, has not been investigated. Here we apply ZooMS screening of Châtelperronian and Early Holocene bone specimens from Quinçay, France, to explore temporal and spatial variation in glutamine deamidation. Our results indicate that chronological resolution is low, while spatial variation is high. Nevertheless, our analysis allows the identification of bone specimens that have undergone diagenetic histories remarkably different (either in length or in type) from spatially related bone specimens. Therefore, ZooMS ammonium-bicarbonate screening is capable of testing bone assemblage homogeneity, which could guide subsequent analysis and interpretation.

Early modern humans (EMH) are often touted as behaviorally advanced to Neandertals, with more sophisticated technologies, expanded resource exploitation, and more complex clothing production. However, recent analyses have indicated that Neandertals were more nuanced in their behavioral adaptations, with the production of the Châtelperronian technocomplex, the processing and cooking of plant foods, and differences in behavioral adaptations according to habitat. This study adds to this debate by addressing the behavioral strategies of EMH (n = 30) within the context of non-dietary anterior tooth-use behaviors to glean possible differences between them and their Neandertal (n = 45) counterparts. High-resolution casts of permanent anterior teeth were used to collect microwear textures of fossil and comparative bioarchaeological samples using a Sensofar white-light confocal profiler with a 100x objective lens. Labial surfaces were scanned, totaling a work envelope of 204 x 276 μm for each individual. The microwear textures were examined for post-mortem damage and uploaded to SSFA software packages for surface characterization. Statistical analyses were performed to examine differences in central tendencies and distributions of anisotropy and textural fill volume variables among the EMH sample itself by habitat, location, and time interval, and between the EMH and Neandertal samples by habitat and location. Descriptive statistics for the EMH sample were compared to seven bioarchaeological samples (n = 156) that utilized different tooth-use behaviors to better elucidate specific activities that may have been performed by EMH. Results show no significant differences between the means within the EMH sample by habitat, location, or time interval. Furthermore, there are no significant differences found here between EMH and Neandertals. Comparisons to the bioarchaeological samples suggest both fossil groups participated in clamping and grasping activities. These results indicate that EMH and Neandertals were similar in their non-dietary anterior tooth-use behaviors and provide additional evidence for overlapping behavioral strategies employed by these two hominins.

The spectacular royal tomb “Ulaankhermiin Shoroon Bumbagar” was discovered in Bulgan province, Mongolia, in 2011. Excavation of the site revealed its internal structure; a slope of 42 meters in length leading down to the underground mausoleum at a depth of 7 m below the ground. Archaeological investigations provided the site with an independent age control suggesting the construction date of the Royal tomb to the last quarter of the VII century. In this study, we directly date different materials from the site, such as terracotta figurines, mortar and host sediment, using infrared-stimulated (IR50) and postinfrared infrared stimulated (pIRIR) luminescence techniques. The most accurate estimate of 670 ± 70AD and 550 ± 110AD was obtained for terracotta figurines using IR50 and pIRIR on aliquots of 4–11 µm polymineral grains. By comparison, sand-sized quartz single grain measurements on the embedded sediment yielded normal equivalent dose (De) distributions with a few outliers, justifying the use of the central age model (CAM) for age calculation, and yielded construction dating to the 780 ± 140AD. Finally, sand-sized quartz single grain measurements on wall mortar revealed incomplete bleaching of grains, requiring detailed analysis using statistical approaches; from this we identified the most well bleached population age of 740 ± 130AD, using the lowest 5%. The luminescence ages are in general agreement with the historically expected age. Statement of significance The archaeological site at Ulaankhermiin Shoroon Bumbagar, Mongolia (with an independent age control) offers a great opportunity to apply luminescence dating methods, such as pIRIR on 4–11 µm polymineral fine grains, and single grain measurements on 200–250 µm quartz grains, to various materials, such as terracotta figurines, mortar-like limestone which covered the walls and embedded sediment in order to test the accuracy of optically stimulated luminescence (OSL) dating techniques. Compared to all other datable archaeological materials, mortar has the advantage of being intentionally manufactured at each stage of construction, since it cannot be recycled. However, dating of the well bleached grains of mortar, which provided the greatest challenge in this study, was achieved using the lowest 5% estimate, yielding an age of 740 ± 130AD. pIRIR on heated polymineral (feldspar-bearing) fine grain aliquots from terracotta figurines was tested, yielding IR50 and pIRIR180 dates in agreement with the CAM estimate from the host sediment. Cite this article Solongo, S., Ochir, A., Tengis, S., Fitzsimmons, K., and Hublin, J.-J. Luminescence dating of mortar and terracotta from a Royal Tomb at Ulaankhermiin Shoroon Bumbagar, Mongolia. STAR 2015; 1(2), STAR2015122054892315Y.0000000004 Data availability The authors confirm that all data underlying the findings are fully available without restriction. All relevant data are contained within the paper.

That great ape endocranial shape development persists into adolescence indicates that the splanchnocranium succeeds brain growth in driving endocranial development. However, the extent of this splanchnocranial influence is unknown. We applied two-block partial least squares analyses of Procrustes shape variables on an ontogenetic series of great ape crania to explore the covariation of the endocranium (the internal braincase) and splanchnocranium (face, or viscerocranium). We hypothesized that a transition between brain growth and splanchnocranial development in the establishment of final endocranial form would be manifest as a change in the pattern of shape covariation between early and adolescent ontogeny. Our results revealed a strong pattern of covariation between endocranium and splanchnocranium, indicating that chimpanzees, gorillas, and orangutans share a common tempo and mode of morphological integration from the eruption of the deciduous dentition onwards to adulthood: a reflection of elongating endocranial shape and continuing splanchnocranial prognathism. Within this overarching pattern, we noted that species variation exists in magnitude and direction, and that the covariation between the splanchnocranium and endocranium is somewhat weaker in early infancy compared to successive age groups. When correcting our covariation analyses for allometry, we found that an ontogenetic signal remains, signifying that allometric variation alone is insufficient to account for all endocranial-splanchnocranial developmental integration. Finally, we assessed the influence of the cranial base, which acts as the interface between the face and endocranium, on the shape of the vault using thin-plate spline warping. We found that not all splanchnocranial shape changes during development are tightly integrated with endocranial shape. This suggests that while the developmental expansion of the brain is the main driver of endocranial shape during early ontogeny, endocranial development from infancy onwards is moulded by the splanchnocranium in conjunction with the neurocranium.

Archaeological sites in northern Africa provide a rich record of increasing importance for the origins of modern human behaviour and for understanding human dispersal out of Africa. However, the timing and nature of Palaeolithic human behaviour and dispersal across north-western Africa (the Maghreb), and their relationship to local environmental conditions, remain poorly understood. The cave of Rhafas (northeast Morocco) provides valuable chronological information about cultural changes in the Maghreb during the Palaeolithic due to its long stratified archaeological sequence comprising Middle Stone Age (MSA), Later Stone Age (LSA) and Neolithic occupation layers. In this study, we apply optically stimulated luminescence (OSL) dating on sand-sized quartz grains to the cave deposits of Rhafas, as well as to a recently excavated section on the terrace in front of the cave entrance. We hereby provide a revised chronostratigraphy for the archaeological sequence at the site. We combine these results with geological and sedimentological multi-proxy investigations to gain insights into site formation processes and the palaeoenvironmental record of the region. The older sedimentological units at Rhafas were deposited between 135 ka and 57 ka (MIS 6 -MIS 3) and are associated with the MSA technocomplex. Tanged pieces start to occur in the archaeological layers around 109 ka, which is consistent with previously published chronological data from the Maghreb. A well indurated duricrust indicates favourable climatic conditions for the pedogenic cementation by carbonates of sediment layers at the site after 57 ka. Overlying deposits attributed to the LSA technocomplex yield ages of ~21 ka and ~15 ka, corresponding to the last glacial period, and fall well within the previously established occupation phase in the Maghreb. The last occupation phase at Rhafas took place during the Neolithic and is dated to ~7.8 ka.

The Neandertal lineage developed successfully throughout western Eurasia and effectively survived the harsh and severely changing environments of the alternating glacial/interglacial cycles from the middle of the Pleistocene until Marine Isotope Stage 3. Yet, towards the end of this stage, at the time of deteriorating climatic conditions that eventually led to the Last Glacial Maximum, and soon after modern humans entered western Eurasia, the Neandertals disappeared. Western Eurasia was by then exclusively occupied by modern humans. We use occlusal molar microwear texture analysis to examine aspects of diet in western Eurasian Paleolithic hominins in relation to fluctuations in food supplies that resulted from the oscillating climatic conditions of the Pleistocene. There is demonstrable evidence for differences in behavior that distinguish Upper Paleolithic humans from members of the Neandertal lineage. Specifically, whereas the Neandertals altered their diets in response to changing paleoecological conditions, the diets of Upper Paleolithic humans seem to have been less affected by slight changes in vegetation/climatic conditions but were linked to changes in their technological complexes. The results of this study also indicate differences in resource exploitation strategies between these two hominin groups. We argue that these differences in subsistence strategies, if they had already been established at the time of the first contact between these two hominin taxa, may have given modern humans an advantage over the Neandertals, and may have contributed to the persistence of our species despite habitat-related changes in food availabilities associated with climate fluctuations.

In many Middle Pleistocene sites, the co-occurrence of hominins with carnivores, who both contributed to faunal accumulations, suggests competition for resources as well as for living spaces. Despite this, there is very little evidence of direct interaction between them to-date. Recently, a human femoral diaphysis has been recognized in South-West of Casablanca (Morocco), in the locality called Thomas Quarry I. This site is famous for its Middle Pleistocene fossil hominins considered representatives of Homo rhodesiensis. The bone was discovered in Unit 4 of the Grotte à Hominidés (GH), dated to c. 500 ky and was associated with Acheulean artefacts and a rich mammalian fauna. Anatomically, it fits well within the group of known early Middle Pleistocene Homo, but its chief point of interest is that the diaphyseal ends display numerous tooth marks showing that it had been consumed shortly after death by a large carnivore, probably a hyena. This bone represents the first evidence of consumption of human remains by carnivores in the cave. Whether predated or scavenged, this chewed femur indicates that humans were a resource for carnivores, underlining their close relationships during the Middle Pleistocene in Atlantic Morocco.

Ochre is the common archaeological term for prehistoric pigments. It is applied to a range of uses, from ritual burials to cave art to medications. While a substantial number of Palaeolithic paint mining pits have been identified across Europe, the link between ochre use and provenance, and their antiquity, has never yet been identified. Here we characterise the mineralogical signature of core-shell processed ochre from the Palaeolithic paint mining pits near Lovas in Hungary, using a novel integration of petrographic and mineralogical techniques. We present the first evidence for core-shell processed, natural pigment that was prepared by prehistoric people from hematitic red ochre. This involved combining the darker red outer shell with the less intensely coloured core to efficiently produce an economical, yet still strongly coloured, paint. We demonstrate the antiquity of the site as having operated between 14-13 kcal BP, during the Epigravettian period. This is based on new radiocarbon dating of bone artefacts associated with the quarry site. The dating results indicate the site to be the oldest known evidence for core-shell pigment processing. We show that the ochre mined at Lovas was exported from the site based on its characteristic signature at other archaeological sites in the region. Our discovery not only provides a methodological framework for future characterisation of ochre pigments, but also provides the earliest known evidence for "value-adding" of products for trade.

For the last ten millennia humans have learned how to manage and produce their own food resources. In Liguria, the exploitation of the marine and coastal environment as well as the acquisition and use of agriculture and pastoralism at the time of Neolithisation (i.e. 6th millennia BC) are evidenced by both archaeological and palaeoenvironmental studies. However, it is difficult, using these methods, to determine the relative contributions to human diets made by foods from these difference sources, especially throughout the Neolithic period. The presence of human burials and animal remains from various sites from this region allows the use of isotopic methods, and in particular the study of the δ13C and δ15N values of bone collagen, which provides information on the sources of dietary protein. This method is able to determine the relative importance of marine foods in the diets of the early Neolithic agriculturalists and pastoralists, as well as indicate dietary differences between contemporary populations. Two coastal Neolithic sites were the subject of our study: Pendimoun (France) and Arene Candide (Italy). The results show, contrary to expectations, the absence of regular consumption of marine resources and the food differences within the populations.

In the northern Adriatic regions, which include the Venetian region and the Dalmatian coast, late Neanderthal settlements are recorded in few sites and even more ephemeral are remains of the Mid-Upper Palaeolithic occupations. A contribution to reconstruct the human presence during this time range has been produced from a recently investigated cave, Rio Secco, located in the northern Adriatic region at the foot of the Carnic Pre-Alps. Chronometric data make Rio Secco a key site in the context of recording occupation by late Neanderthals and regarding the diffusion of the Mid-Upper Palaeolithic culture in a particular district at the border of the alpine region. As for the Gravettian, its diffusion in Italy is a subject of on-going research and the aim of this paper is to provide new information on the timing of this process in Italy. In the southern end of the Peninsula the first occupation dates to around 28,000 14C BP, whereas our results on Gravettian layer range from 29,390 to 28,995 14C years BP. At the present state of knowledge, the emergence of the Gravettian in eastern Italy is contemporaneous with several sites in Central Europe and the chronological dates support the hypothesis that the Swabian Gravettian probably dispersed from eastern Austria.

The dental human remains from the early Aurignacian layers of Brassempouy (Landes) have been recently described by Henry-Gambier et al. (2004). We provide a critical re-assessment of the features that have led these authors to conclude that the taxonomic status of these fossils is uncertain. Although the works of one of us (S.B.) have been partly used and cited by Henry-Gambier et al. (2004), we disagree with the conclusions that have been drawn from them. In our view and based on the available evidence, the early Aurignacian dental remains from Brassempouy are unambiguously modern in their anatomy. They indeed provide further evidence that the makers of the ancient Aurignacian were early anatomically modern Europeans.