The use of bones as tools in Late Lower Paleolithic of Central Italy.

The Latium area in Italy has yielded rich evidence of Lower Paleolithic sites with both faunal remains, artefacts, and human fossil remains, such as the Ceprano human skull. Many are the sites where lithic industry has been found in association with bone industry. Medium and large animals were a key resource because they provided an enormous amount of meat and fat. However, they were extensively exploited for their bones, rich in marrow, and as raw material for tool production. Bone tools are so far few documented for early period of time and especially for the Middle Pleistocene in Western Europe. We report here evidence of bone tools and their efficiency of use for hominin groups living in the Frosinone-Ceprano basin during the MIS 11/10, a key period which records behavioral innovations and onset of the Neanderthal behaviors. In three sites, Isoletta, Colle Avarone and Selvotta, several bone tools and bone flakes have been discovered (MIS 11/10). They were associated to stone artefacts part of the hominins tool-kit. Technological and use-wear analyses conducted on these bone industries, dated between 410 and 430 ka, yield relevant results to understand the effectiveness of the bones tools found associated with lithic series, including handaxes.

With Impressions Chosen from Another Time: Core Technologies and Debitage Production at the Lower Palaeolithic Site of Notarchirico (670-695 ka; layers F to I2).

The earliest evidence of bifaces in western Europe is dated to the initial phase of the Middle Pleistocene (la Noira, Notarchirico, Moulin Quignon, 700-670 ka), with the findings of Barranc de la Boella (1.0-0.9 Ma) considered to be an earlier local evolution. No transition assemblages are recorded during this time frame, and the "abrupt" appearance of bifaces during this time frame is associated with significant cognitive shifts in human technological behaviours (Acheulean techno-complex). The new investigations conducted at the site of Notarchirico unearthed 30 ka of repeated human occupation (695-670 ka, layers F-I2) during MIS 17, with evidence of bifacial tools in layer G (680 ka) and F along with other heavy-duty implements (LCTs, pebble tools, etc.). Massive production of debitage products realised on local raw materials collected in situ through simple and efficient core technologies characterises a large part of the lithic assemblage with a high ratio of diversified light-duty tools, including modified chert nodules. Despite core and flake assemblages being a recurrent trait of Lower Pleistocene contexts, the increase in retouched implements recorded at the onset of the Middle Pleistocene has been considered a significant technological shift. The technological analysis of the debitage products presented in this work highlights recurrent and systematic technological behaviours of the hominins of Notarchirico-who proved to efficiently overcome the raw materials dimensional constraints-even in the layers without bifaces. This may shed light on the meaning of cultural and behavioural innovation that the Acheulean techno-complex is thought to bring over Europe. It is plausible that given the substantial homogeneity of the lithic strategies within the sequence of Notarchirico, which only the "introduction" of the bifaces in the upper layers seems to interrupt, a supposed behavioural or cultural change in the site might have already occurred in the lowermost portion of the sequence. In this work, we evaluate the degree of change-if any-from a technological perspective by analysing the debitage reduction sequences.

The Western European Acheulean: Reading variability at a regional scale.

In the context of the Western European Acheulean Project, this study aims to characterize Acheulean technology in Western Europe through the analysis of handaxes and cleavers from 10 key sites (Britain 4, France 4, and Spain 2) to acquire a regional view of the occupation. The historically different systems used to categorize and analyze the data have made it difficult to compare results. Here we apply a unified and simple method (Western European Acheulean Project) that combines the traditional technological and metrical analysis of assemblages containing handaxes and cleavers with an in-depth geometric morphometric approach using three-dimensional models. This approach allows us to achieve a regional interpretation that identifies innovations through time and shaping strategies across the area. Our findings indicate the existence of two main technological groups in the sampled record: 1) northwestern and central France and Britain, from MIS 17/16 to MIS 11, and 2) Atlantic edge (Atapuerca in Spain and Menez-Dregan in France), from MIS 12/11 to MIS 8. Based on our technological analysis, the shaping of handaxes and cleavers was developed through time as a continuum of accumulative actions, with longer and more complex shaping strategies over time. Shaping technology shows traditions of manufacture over both time and geographical areas, which suggest cultural diffusion. Our geometric morphometric analysis further helped to identify not only general trends but also local adaptations in handaxe forms. Based on our findings, there were no apparent sudden innovations, but rather the application and development of specific techniques to refine size and shape.

Multi-isotope zooarchaeological investigations at Abri du Maras: The paleoecological and paleoenvironmental context of Neanderthal subsistence strategies in the Rhône Valley during MIS 3.

The exploitation of mid- and large-sized herbivores (ungulates) was central to hominin subsistence across Late Pleistocene Europe. Reconstructing the paleoecology of prey-taxa is key to better understanding procurement strategies, decisions and behaviors, and the isotope analysis of faunal bones and teeth found at archaeological sites represent a powerful means of accessing information about past faunal behaviors. These isotope zooarchaeological approaches also have a near-unique ability to reveal environmental conditions contemporary to the human activities that produced these remains. Here, we present the results of a multi-isotope, multitissue study of ungulate remains from the Middle Paleolithic site of Abri du Maras, southern France, providing new insights into the living landscapes of the Rhône Valley during MIS 3 (level 4.2 = 55 ± 2 to 42 ± 3 ka; level 4.1 = 46 ± 3 to 40 ± 3 ka). Isotope data (carbon, nitrogen) reveal the dietary niches of different ungulate taxa, including the now-extinct giant deer (Megaloceros). Oxygen isotope data are consistent with a mild seasonal climate during level 4.2, where horse (Equus), bison (Bison), and red deer (Cervus elaphus) were exploited year-round. Strontium and sulfur isotope analyses provide new evidence for behavioral plasticity in Late Pleistocene European reindeer (Rangifer) between level 4.2 and level 4.1, indicating a change from the migratory to the sedentary ecotype. In level 4.1, the strong seasonal nature of reindeer exploitation, combined with their nonmigratory behavior, is consistent with a seasonally restricted use of the site by Neanderthals at that time or the preferential hunting of reindeer when in peak physical condition during the autumn.

New evidence of Neandertal butchery traditions through the marrow extraction in southwestern Europe (MIS 5-3).

Long bone breakage for bone marrow recovery is a commonly observed practice in Middle Palaeolithic contexts, regardless of the climatic conditions. While lithic technology is largely used to define cultural patterns in human groups, despite dedicating research by zooarchaeologists, for now butchering techniques rarely allowed the identification of clear traditions, notably for ancient Palaeolithic periods. In this paper, we test the hypothesis of butchery traditions among Neandertal groupsusing the bone assemblages from three sites in southwestern Europe. These sites are located in southeastern France and northern Italy and are dated to the Late Middle Palaeolithic: Abri du Maras (Marine Isotopic Stages (MIS) 4-3, Ardèche), Saint-Marcel (MIS 3, Ardèche), and Riparo Tagliente (MIS 4-3, Verona). The detection of culturally-induced patterns of bone breakage involves differentiating them from intuitively generated patterns. To tackle this issue, we used a zooarchaeological approach focusing on the percussion marks produced during the bone breakage process. Statistical analyses as the chi-square test of independence were employed to verify if percussion mark locations were randomly distributed, and if these distributions were different from the intuitive ones. For femurs and humeri, our results demonstrate that Neandertal groups occupying the Abri du Maras (levels 4.1 and 4.2) and the Saint-Marcel Cave (levels g and h) sites in France applied butchery traditions to recover yellow marrow. However, the traditions developed at each site were different. On the contrary, in Riparo Tagliente, in Italy, several groups or individuals of a same group did not share the same butchery traditions over time. Regarding the Abri du Maras and Saint Marcel Cave assemblages, our research demonstrates that Neandertal groups applied intense standardized bone breakage, far from the intuitive practice observed experimentally and related to bone density and/or skeletal morphology. These standardized patterns, which are systematic and counter-intuitive, can be interpreted as culturally induced for the Abri du Maras and Saint Marcel Cave. The diversity of Neandertal traditions should be considered by taking into account the butchery, in particular the practice of bone marrow extraction, and not only technological behaviours and types of tool kits.

Palaeolithic polyhedrons, spheroids and bolas over time and space.

Polyhedrons, spheroids and bolas (PSBs) are present in lithic series from the Lower Palaeolithic onwards and are found in several regions of the world. Nevertheless, very little is known about them. We propose here to summarise, illustrate and discuss the current state of our knowledge about these artefacts. Based on the available data in the literature and on our observations of several collections, we set up a database comprising 169 Palaeolithic assemblages with PSBs. Thanks to the statistical analysis of these data, we aim to highlight potential relationships between PSB characteristics (e.g., quantity, raw material) and assemblage composition and context, according to regions and chrono-cultural attributions. We also aim to discuss the question of artefacts from possible independent local histories, especially in Northwest Europe, where these objects are scarce. Our study concludes that hard stones (stones with high resistance to a physical constraint) available locally were generally selected to produce PSBs. Soft sedimentary rocks are suitable for their manufacture, and were selected too, whereas siliceous materials were left aside. We hypothesise that the scarcity of PSBs in Northwest Europe could result from a combination of cultural and environmental factors: it could be part of a regional tradition, influenced by the abundance of siliceous materials in the environment. In this region where the lithic production is widely made of flint, even though other materials were available, objects made from hard stones are scarce, resulting in a toolkit with only rare PSBs and cleavers. Was flint too brittle for the functions of PSBs? Raw materials of PSBs are often similar to those of heavy-duty tools in assemblages, which could provide other clues about their functions (e.g., tasks requiring a resistance to shocks). It is possible that their raw materials partly conditioned their final shape. PSBs can comprise a wide variety of artefacts, that for some could have change of status (e.g., from cores to percussive tools), diffused, adapted but also reinvented over two million years.

Reappraisal of the chronology of Orgnac 3 Lower-to-Middle Paleolithic site (Ardèche, France), a regional key sequence for the Middle Pleistocene of southern France.

Previous studies have suggested that the Lower-to-Middle Paleolithic transition was associated with the earliest Neanderthals, but recent research has established that the oldest Neanderthal fossils and the first signs of their technologies and behavior appear from MIS 11 or possibly earlier. To understand these changes, re-evaluation of the evidence is necessary to determine if this transition corresponds to a progressive evolution rather than abrupt change. Orgnac 3 is a key and appropriate site to study this research context. Located in southern France, it yields a long stratigraphic sequence testifying the evolution of technical and subsistence behaviors of pre-Neanderthal human groups during a Middle Pleistocene interglacial-glacial cycle. In this article, a new chronological framework is provided for the sequence based on results of dating methods applied to various types of geological materials. Speleothems and volcanic minerals, dated in previous studies by U-series and 40Ar/39Ar, respectively, show periods of calcitic crystallization and regional volcanic activity. Other materials, such as heated flints and herbivore teeth, are directly related to evidence of anthropogenic activities and are analyzed in the present work by trapped-charge dating methods such as thermoluminescence and electron spin resonance combined with uranium series (ESR/U-series). The new thermoluminescence and ESR/U-series dates confirm the attribution of the Orgnac 3 stratigraphic sequence to the MIS 10-MIS 8 period and are discussed in relation to paleoenvironmental data derived from bioarchaeological studies. The paleoanthropological levels, including the emergence of Levallois technology, are dated to ca. 275 ka (early MIS 8) and appear coeval to a wet and temperate period recorded locally, the Amargiers interstadial, defined in the regional palynological records. The implications of this reassessed chronology for the archaeological assemblages are discussed in the wider context of behavioral innovations from MIS 11 onward and their establishment in subsequent periods.

Acheulean variability in Western Europe: The case of Menez-Dregan I (Plouhinec, Finistère, France).

The establishment of the Acheulean in Europe occurred after MIS 17, but it was after the harsh glaciation of MIS 12 and during the long interglacial of MIS 11 that human occupation of Western Europe became more sustained, with an increased number of sites. Menez-Dregan I (Brittany, France) is one of the key sites in Western Europe that dates from this threshold, with an alternating sequence of 16 occupation levels and four marine deposits, from MIS 12 to 8. The large lithic assemblages of more than 154,000 artifacts from knapping (cores, flakes) and shaping (macrotools and shaping flakes) show the varying use of raw materials and activities at the site through the sequence. This work focuses on the study of the handaxes and cleavers using technological and metrical methods with multivariate analysis, in combination with geometric morphometrics, and places these analyses within the context of other technological changes at the site. Collectively, results show the persistent use through the sequence of the same lithic raw materials and technologies, including fire use and the import of glossy sandstone from 20 km away, but with variation in activities at the site. These findings suggest that Menez-Dregan I shows the development of a specific material culture that reflects the local resources and environment. Results further indicate that the site shows the sustained hominin occupation of the area, despite varying climate and environment, with strong traditions of social learning that were maintained through flexibility of site use, deep understanding of the local territory, and the innovation of new technologies, such as the use of fire. Evidence from the site is placed within the wider context of Europe, and contrasted with areas to the north, such as Britain, where hominin occupation was more sporadic and driven by cyclical climate change.

A way to break bones? The weight of intuitiveness.

During the Paleolithic period, bone marrow extraction was an essential source of fat nutrients for hunter-gatherers especially throughout cold and dry seasons. This is attested by the recurrent findings of percussion marks in osteological material from anthropized archaeological levels. Among them some showed indicators that the marrow extraction process was part of a butchery cultural practice, meaning that the inflicted fracturing gestures and techniques were recurrent, standardized and counter-intuitive. In order to assess the weight of the counter-intuitive factor in the percussion mark pattern distribution, we carried out an experiment that by contrast focuses on the intuitive approach of fracturing bones to extract marrow, involving individual without experience in this activity. We wanted to evaluate the influence of bone morphology and the individuals' behaviour on the distribution of percussion marks. Twelve experimenters broke 120 limb bones, a series of 10 bones per individual. During the experiment, information concerning the fracture of the bones as well as individual behaviour was collected and was subsequently compared to data from the laboratory study of the remains. Then, we applied an innovative GIS (Geographic Information System) method to analyze the distribution of percussion marks to highlight recurrent patterns. Results show that in spite of all the variables there is a high similarity in the distribution of percussion marks which we consider as intuitive patterns. The factor influenced the distribution for the humerus, radius-ulna and tibia series is the bone morphology, while for the femur series individual behaviour seems to have more weight in the distribution. To go further in the subject we need to compare the intuitive models with the distributions of percussion marks registered in fossil assemblages. Thus, it would be possible to propose new hypotheses on butchering practices based on the results presented in this work.

Tracking behavioral persistence and innovations during the Middle Pleistocene in Western Europe. Shift in occupations between 700 and 450 ka at la Noira site (Centre, France).

Some areas in Western Europe indicate hiatuses in human occupations, which cannot be systematically attributed to taphonomic factors and poor site preservation. The site of la Noira in the center of France records two occupation phases with a significant time gap. The older one is dated to around 700 ka (stratum a) with an Acheulean assemblage, among the earliest in Western Europe, and the upper phase of the sequence (stratum c) is dated to ca. 450 ka. Humans left the area at around 670 ka, at the beginning of the marine isotope stage (MIS) 16 glacial stage, when cold conditions became too severe. No sites between 650 and 450 ka have yet been discovered in the center region despite systematic surveys over the past three decades. The archaeological evidence indicates that populations returned to the area, at the end of MIS 12 or the beginning of the long interglacial MIS 11. Here, we use technological behaviors common to the two levels of la Noira-strata a and c to evaluate their differences. Compared to other key European sequences, this site can be used to address the evolution of the behavioral strategies in Europe between MIS 17 and 11. We formulate two hypotheses concerning the human settlement of this area: (1) local behavioral evolution over time of populations occasionally occupying the region when the climate was favorable or (2) dispersal and arrival of new populations from other areas. The results focus on (1) changes in land-use patterns with the extension of the territory used by hominins in the upper level, (2) the introduction of new core technologies, including some evidence of early Levallois debitage, and (3) more intensive shaping of bifaces and bifacial tools. Results attest that the la Noira archaeological assemblages record similar regional behavioral evolution as observed at a larger scale in Europe.

First 40Ar/39Ar analyses of Australasian tektites in close association with bifacially worked artifacts at Nalai site in Bose Basin, South China: The question of the early Chinese Acheulean.

The recently discovered Nalai site is one of the Bose Basin localities, which is key to studying the earliest bifaces in China. The Nalai site has yielded an abundance of lithic artifacts, including bifaces and tektites in close association. The total fusion 40Ar/39Ar method was applied to four tektites discovered beside and contemporaneous with bifaces in the red laterite sediments of the upper levels of the T4 terrace (layers 4 and 5). Our 40Ar/39Ar data with a weighted mean age of 809 ± 12 ka provide for the first time unequivocal dates for bifacial production at Bose, broadly consistent with the precise Australasian tektite age of 788.1 ± 2.8 ka, recently published by other investigators. The relatively important errors reported here suggest sample contamination by clasts or bubbles for the oldest aliquots and alteration for the younger ones. The lithic assemblage from layers 4 and 5 of the Nalai site is quite similar to that found at other sites in the Bose Basin. The assemblages are dominated by choppers, but bifaces, picks, and unifaces give a Mode 2 and Acheulean-type character to the series. The high frequency of the round tongue-shaped tip, a low elongation index, and a wide and thick base characterize the Large Cutting Tools. These results contribute to resolving ongoing debates on the timing and origin of bifaces and the Acheulean in China.

An ecological niche shift for Neanderthal populations in Western Europe 70,000 years 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.

The implications of thumb movements for Neanderthal and modern human manipulation.

Much research has debated the technological abilities of Neanderthals relative to those of early modern humans, with a particular focus on subtle differences in thumb morphology and how this may reflect differences in manipulative behaviors in these two species. Here, we provide a novel perspective on this debate through a 3D geometric morphometric analysis of shape covariation between the trapezial and proximal first metacarpal articular surfaces of Neanderthals (Homo neanderthalensis) in comparison to early and recent humans (Homo sapiens). Results show a distinct pattern of shape covariation in Neanderthals, consistent with more extended and adducted thumb postures that may reflect habitual use of grips commonly used for hafted tools. Both Neanderthals and recent humans demonstrate high intraspecific variation in shape covariation. This intraspecific variation is likely the result of genetic and/or developmental differences, but may also reflect, in part, differing functional requirements imposed by the use of varied tool-kits. These results underscore the importance of holistic joint shape analysis for understanding the functional capabilities and evolution of the modern human thumb.

The origin of early Acheulean expansion in Europe 700 ka ago: new findings at Notarchirico (Italy).

Notarchirico (Southern Italy) has yielded the earliest evidence of Acheulean settlement in Italy and four older occupation levels have recently been unearthed, including one with bifaces, extending the roots of the Acheulean in Italy even further back in time. New 40Ar/39Ar on tephras and ESR dates on bleached quartz securely and accurately place these occupations between 695 and 670 ka (MIS 17), penecontemporaneous with the Moulin-Quignon and la Noira sites (France). These new data demonstrate a very rapid expansion of shared traditions over Western Europe during a period of highly variable climatic conditions, including interglacial and glacial episodes, between 670 and 650 (i.e., MIS17/MIS16 transition). The diversity of tools and activities observed in these three sites shows that Western Europe was populated by adaptable hominins during this time. These conclusions question the existence of refuge areas during intense glacial stages and raise questions concerning understudied migration pathways, such as the Sicilian route.

The Middle Pleistocene hominin mandible from Payre (Ardèche, France).

Although Neandertals are the best-known fossil hominins, the tempo and evolutionary processes in their lineage are strongly debated. This is in part due to the scarcity of the fossil record, in particular before the marine isotopic stage (MIS) 5. In 2010, a partial hominin mandible was discovered at the Middle Paleolithic site of Payre (France) in a layer that is dated to the end of MIS 8/beginning of MIS 7, a time period for which very few fossils are known in Europe. The Payre 15 mandible retains the complete symphyseal region and right lateral corpus with heavily worn P4, M1, and M2 in situ. Taphonomic modifications in the form of three notches suggest that this individual was chewed by a carnivore. We provide here the first detailed description of this specimen and a comparative analysis that includes morphological features, linear mandibular dimensions, an elliptic Fourier analysis of the symphysis, and a morphometric analysis of the M1 roots (based on segmented CT scan data). Our comparative sample encompasses European Middle and Upper Pleistocene specimens attributed to Homo heidelbergensis and Homo neanderthalensis, Upper Pleistocene Homo sapiens, and Holocene Homo sapiens. The Payre 15 mandible shows a combination of primitive and Neandertal-like features, with a receding symphyseal profile without any element of the mentum osseum, a posterior location of the mental foramen and lateral prominence. Its mandibular body is tall and thick anteriorly. Payre 15 has mesotaurodont M1 roots and a three-rooted M2. By its dimensions and combination of features, Payre 15 aligns better with Middle Pleistocene European hominins than with MIS 6-3 Neandertals. Noteworthy, it falls well within the range of variation of the Sima de los Huesos sample. Our results underscore that the total pattern of Neandertal-derived morphology was not achieved at the beginning of the MIS 7 and suggest a low level of mandibular diachronic changes for the period MIS 11-7.

Early Levallois core technology between Marine Isotope Stage 12 and 9 in Western Europe.

Early Levallois core technology is usually dated in Europe to the end of Marine Isotope Stage (MIS) 9 and particularly from the beginning of MIS 8 to MIS 6. This technology is considered as one of the markers of the transition from lower to Middle Paleolithic or from Mode 2 to Mode 3. Recent discoveries show that some lithic innovations actually appeared earlier in western Europe, from MIS 12 to MIS 9, contemporaneous with changes in subsistence strategies and the first appearance of early Neanderthal anatomical features. Among these discoveries, there is the iconic Levallois core technology. A selection of well-dated assemblages in the United Kingdom, France, and Italy dated from MIS 12 to 9, which include both cores and flakes with Levallois features, has been described and compared with the aim of characterizing this technology. The conclusion supports the interpretation that several technical features may be attributed to a Levallois technology similar to those observed in younger Middle Paleolithic sites, distinct from the main associated core technologies in each level. Some features in the sample of sites suggest a gradual transformation of existing core technologies. The small evidence of Levallois could indicate occasional local innovations from different technological backgrounds and would explain the diversity of Levallois methods that is observed from MIS 12. The technological roots of Levallois technology in the Middle Pleistocene would suggest a multiregional origin and diffusion in Europe and early evidence of regionalization of local traditions through Europe from MIS 12 to 9. The relationships of Levallois technology with new needs and behaviors are discussed, such as flake preference, functional reasons related to hunting and hafting, an increase in the use of mental templates in European populations, and changes in the structure of hominin groups adapting to climatic and environmental changes.

Correction: A biface production older than 600 ka ago at Notarchirico (Southern Italy) contribution to understanding early Acheulean cognition and skills in Europe.

[This corrects the article DOI: 10.1371/journal.pone.0218591.].

The earliest evidence of Acheulian occupation in Northwest Europe and the rediscovery of the Moulin Quignon site, Somme valley, France.

The dispersal of hominin groups with an Acheulian technology and associated bifacial tools into northern latitudes is central to the debate over the timing of the oldest human occupation of Europe. New evidence resulting from the rediscovery and the dating of the historic site of Moulin Quignon demonstrates that the first Acheulian occupation north of 50°N occurred around 670-650 ka ago. The new archaeological assemblage was discovered in a sequence of fluvial sands and gravels overlying the chalk bedrock at a relative height of 40 m above the present-day maximal incision of the Somme River and dated by ESR on quartz to early MIS 16. More than 260 flint artefacts were recovered, including large flakes, cores and five bifaces. This discovery pushes back the age of the oldest Acheulian occupation of north-western Europe by more than 100 ka and bridges the gap between the archaeological records of northern France and England. It also challenges hominin dispersal models in Europe showing that hominins using bifacial technology, such as Homo heidelbergensis, were probably able to overcome cold climate conditions as early as 670-650 ka ago and reasserts the importance of the Somme valley, where Prehistory was born at the end of the 19th century.