Correction: Elephant bones for the Middle Pleistocene toolmaker.

[This corrects the article DOI: 10.1371/journal.pone.0256090.].

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.

A marine isotope stage 11 coastal Acheulian workshop with associated wood at Amanzi Springs Area 1, South Africa.

Amanzi Springs is a series of inactive thermal springs located near Kariega in the Eastern Cape of South Africa. Excavations in the 1960s exposed rare, stratified Acheulian-bearing deposits that were not further investigated over the next 50 years. Reanalysis of the site and its legacy collection has led to a redefined stratigraphic context for the archaeology, a confirmed direct association between Acheulian artefacts and wood, as well as the first reliable age estimates for the site. Thermally transferred optically stimulated luminescence and post-infrared infrared stimulated luminescence dating indicates that the Acheulian deposits from the Amanzi Springs Area 1 spring eye formed during Marine Isotope Stage (MIS) 11 at ~ 404-390 ka. At this time, higher sea levels of ~13-14m would have placed Amanzi Springs around 7 km from a ria that would have formed along what is today the Swartkops River, and which likely led to spring reactivation. This makes the Amanzi Springs Area 1 assemblage an unusual occurrence of a verified late occurring, seaward, open-air Acheulian occupation. The Acheulian levels do not contain any Middle Stone Age (MSA) elements such as blades and points that have been documented in the interior of South Africa at this time. However, a small number of stone tools from the upper layers of the artefact zone, and originally thought of as intrusive, have been dated to ~190 ka, at the transition between MIS 7 to 6, and represent the first potential MSA identified at the site.

A refined chronology for the Middle and early Upper Paleolithic sequence of Riparo Mochi (Liguria, Italy).

The Riparo Mochi rock shelter, located on the Ligurian coast of Italy, is one of the most important early Upper Paleolithic sites on the Mediterranean rim. Its ∼10-m-deep stratigraphy comprises a Mousterian sequence, followed by various development stages of the Upper Paleolithic. A series of radiometric dates on marine shells bearing traces of human modification has provided a chronological framework for the final Mousterian and the Proto-Aurignacian of the site. Based on modeling results, the end of the Mousterian was dated between 44.0 and 41.8 ka cal BP (68% probability) and the beginning of the Proto-Aurignacian between 42.7 and 41.6 ka cal BP (68% probability). However, these estimates were based on a limited number of radiocarbon ages in the Mousterian levels. Here, we report new dating of the Mochi sequence using luminescence techniques, along with new radiocarbon measurements. The combination of these results using a Bayesian modeling approach allows for the first time the establishment of a more precise timing for the Mousterian occupation at the site. We show that Mousterian groups were already present at Riparo Mochi by at least 65 ka and continued to occupy the site for another 20 ka. The transition to the earliest Upper Paleolithic at the site is centered around 44.3-41.1 ka (95.4% probability), providing our best age estimate for the beginning of the Early Upper Paleolithic and the establishment of modern human groups in the Balzi Rossi. The sequence continues upward with a more evolved Aurignacian phase and a Gravettian phase starting at ∼26 ka or earlier.

The Thorny Issue of African Porcupines: a New Mandible of Hystrix makapanensis from Olduvai Gorge (Tanzania) and Rediagnosis of the Species.

Several porcupine taxa are reported from the middle Miocene to the early Holocene in the Old World. Among these, five species of the subfamily Hystricinae occurred in Africa approximately in the last 6 Ma: the extinct Hystrix makapanensis, Hystrix leakeyi, and Xenohystrix crassidens and the still living Hystrix africaeaustralis and Hystrix cristata. The large-sized H. makapanensis is reported from numerous sites in East and South Africa between the early Pliocene and Early Pleistocene. In this paper, we describe a new mandible of H. makapanensis from the world-renowned Tanzanian paleontological and archeological site of Olduvai Gorge (HWK West; lowermost Bed II; ca. 1.8-1.7 Ma). The discovery of the new mandible triggered a comprehensive review of the entire African record of H. makapanensis. In particular, we describe or re-analyze the samples from South Africa (Makapansgat Limeworks, Gondolin, Kromdraai, Swartkrans, and Sterkfontein), Tanzania (Olduvai and Laetoli), Ethiopia (Omo Shungura and Hadar), and Kenya (Chemeron), enriching the quantity of specimens confidently referable to this species and above all improving the information on its craniodental anatomy. On this basis, we: (1) propose an emended diagnosis of H. makapanensis; (2) point out the morphological and biometric differences between H. makapanensis and other African Hystricinae (also in terms of body mass); and (3) broaden the knowledge on the geographical and chronological distribution of this extinct species. Supplementary Information: The online version contains supplementary material available at 10.1007/s10914-021-09588-z.

Elephant bones for the Middle Pleistocene toolmaker.

The use of bone as raw material for implements is documented since the Early Pleistocene. Throughout the Early and Middle Pleistocene bone tool shaping was done by percussion flaking, the same technique used for knapping stone artifacts, although bone shaping was rare compared to stone tool flaking. Until recently the generally accepted idea was that early bone technology was essentially immediate and expedient, based on single-stage operations, using available bone fragments of large to medium size animals. Only Upper Paleolithic bone tools would involve several stages of manufacture with clear evidence of primary flaking or breaking of bone to produce the kind of fragments required for different kinds of tools. Our technological and taphonomic analysis of the bone assemblage of Castel di Guido, a Middle Pleistocene site in Italy, now dated by 40Ar/39Ar to about 400 ka, shows that this general idea is inexact. In spite of the fact that the number of bone bifaces at the site had been largely overestimated in previous publications, the number of verified, human-made bone tools is 98. This is the highest number of flaked bone tools made by pre-modern hominids published so far. Moreover the Castel di Guido bone assemblage is characterized by systematic production of standardized blanks (elephant diaphysis fragments) and clear diversity of tool types. Bone smoothers and intermediate pieces prove that some features of Aurignacian technology have roots that go beyond the late Mousterian, back to the Middle Pleistocene. Clearly the Castel di Guido hominids had done the first step in the process of increasing complexity of bone technology. We discuss the reasons why this innovation was not developed. The analysis of the lithic industry is done for comparison with the bone industry.

Drimolen cranium DNH 155 documents microevolution in an early hominin species.

Paranthropus robustus is a small-brained extinct hominin from South Africa characterized by derived, robust craniodental morphology. The most complete known skull of this species is DNH 7 from Drimolen Main Quarry, which differs from P. robustus specimens recovered elsewhere in ways attributed to sexual dimorphism. Here, we describe a new fossil specimen from Drimolen Main Quarry, dated from approximately 2.04-1.95 million years ago, that challenges this view. DNH 155 is a well-preserved adult male cranium that shares with DNH 7 a suite of primitive and derived features unlike those seen in adult P. robustus specimens from other chronologically younger deposits. This refutes existing hypotheses linking sexual dimorphism, ontogeny and social behaviour within this taxon, and clarifies hypotheses concerning hominin phylogeny. We document small-scale morphological changes in P. robustus associated with ecological change within a short time frame and restricted geography. This represents the most highly resolved evidence yet of microevolutionary change within an early hominin species.

Contemporaneity of Australopithecus, Paranthropus, and early Homo erectus in South Africa.

Understanding the extinction of Australopithecus and origins of Paranthropus and Homo in South Africa has been hampered by the perceived complex geological context of hominin fossils, poor chronological resolution, and a lack of well-preserved early Homo specimens. We describe, date, and contextualize the discovery of two hominin crania from Drimolen Main Quarry in South Africa. At ~2.04 million to 1.95 million years old, DNH 152 represents the earliest definitive occurrence of Paranthropus robustus, and DNH 134 represents the earliest occurrence of a cranium with clear affinities to Homo erectus These crania also show that Homo, Paranthropus, and Australopithecus were contemporaneous at ~2 million years ago. This high taxonomic diversity is also reflected in non-hominin species and provides evidence of endemic evolution and dispersal during a period of climatic variability.

Complexities of assessing palaeocave stratigraphy: reconstructing site formation of the ∼2.61 Ma Drimolen Makondo fossil site.

Palaeocave sites in South Africa are world renowned repositories for palaeontological and archaeological material, dating from the terminal Pliocene to the Early Pleistocene. Due to their antiquity, complex karstification history and multifaceted infilling phases, palaeocave sites are notoriously difficult to contextualise. Further to this, 19th century lime-mining and diverse excavation and sampling techniques, have complicated stratigraphic interpretations of fossil-bearing deposits within the region. Locating and assessing newly discovered, minimally disturbed palaeocave sites allow for contextual information to be gathered with greater confidence and can aid in constructing a more robust understanding of the South African fossil record. Here, we use Drimolen Makondo; a minimally lime-mined ∼2.61 Ma palaeontological site, to apply a series of in-depth stratigraphic and micromorphological studies. Contextual data presented within this study, testifies to a relatively rapid infill with greater fluvial activity when compared to adjacent deposits at the younger ∼2.04-1.95 Ma Drimolen Main Quarry. The quantity of articulated macromammalian remains, high density of micromammalian remains and pollen identified, also highlights Drimolen Makondo as a key site for ongoing palaeoenvironmental studies at the Pliocene to Pleistocene transition in South Africa.

The Middle Pleistocene (MIS 12) human dental remains from Fontana Ranuccio (Latium) and Visogliano (Friuli-Venezia Giulia), Italy. A comparative high resolution endostructural assessment.

The penecontemporaneous Middle Pleistocene sites of Fontana Ranuccio (Latium) and Visogliano (Friuli-Venezia Giulia), set c. 450 km apart in central and northeastern Italy, respectively, have yielded some among the oldest human fossil remains testifying to a peopling phase of the Italian Peninsula broadly during the glacial MIS 12, a stage associated with one among the harshest climatic conditions in the Northern hemisphere during the entire Quaternary period. Together with the large samples from Atapuerca Sima de los Huesos, Spain, and Caune de l'Arago at Tautavel, France, the remains from Fontana Ranuccio and Visogliano are among the few mid-Middle Pleistocene dental assemblages from Western Europe available for investigating the presence of an early Neanderthal signature in their inner structure. We applied two- three-dimensional techniques of virtual imaging and geometric morphometrics to the high-resolution X-ray microtomography record of the dental remains from these two Italian sites and compared the results to the evidence from a selected number of Pleistocene and extant human specimens/samples from Europe and North Africa. Depending on their preservation quality and on the degree of occlusal wear, we comparatively assessed: (i) the crown enamel and radicular dentine thickness topographic variation of a uniquely represented lower incisor; (ii) the lateral crown tissue proportions of premolars and molars; (iii) the enamel-dentine junction, and (iv) the pulp cavity morphology of all available specimens. Our analyses reveal in both samples a Neanderthal-like inner structural signal, for some aspects also resembling the condition shown by the contemporary assemblage from Atapuerca SH, and clearly distinct from the recent human figures. This study provides additional evidence indicating that an overall Neanderthal morphological dental template was preconfigured in Western Europe at least 430 to 450 ka ago.

The dawn of dentistry in the late upper Paleolithic: An early case of pathological intervention at Riparo Fredian.

OBJECTIVES: Early evidence for the treatment of dental pathology is found primarily among food-producing societies associated with high levels of oral pathology. However, some Late Pleistocene hunter-gatherers show extensive oral pathology, suggesting that experimentation with therapeutic dental interventions may have greater antiquity. Here, we report the second earliest probable evidence for dentistry in a Late Upper Paleolithic hunter-gatherer recovered from Riparo Fredian (Tuscany, Italy). MATERIALS AND METHODS: The Fredian 5 human consists of an associated maxillary anterior dentition with antemortem exposure of both upper first incisor (I1 ) pulp chambers. The pulp chambers present probable antemortem modifications that warrant in-depth analyses and direct dating. Scanning electron microscopy, microCT and residue analyses were used to investigate the purported modifications of external and internal surfaces of each I1 . RESULTS: The direct date places Fredian 5 between 13,000 and 12,740 calendar years ago. Both pulp chambers were circumferentially enlarged prior to the death of this individual. Occlusal dentine flaking on the margin of the cavities and striations on their internal aspects suggest anthropic manipulation. Residue analyses revealed a conglomerate of bitumen, vegetal fibers, and probable hairs adherent to the internal walls of the cavities. DISCUSSION: The results are consistent with tool-assisted manipulation to remove necrotic or infected pulp in vivo and the subsequent use of a composite, organic filling. Fredian 5 confirms the practice of dentistry-specifically, a pathology-induced intervention-among Late Pleistocene hunter-gatherers. As such, it appears that fundamental perceptions of biomedical knowledge and practice were in place long before the socioeconomic changes associated with the transition to food production in the Neolithic.

New footprints from Laetoli (Tanzania) provide evidence for marked body size variation in early hominins.

Laetoli is a well-known palaeontological locality in northern Tanzania whose outstanding record includes the earliest hominin footprints in the world (3.66 million years old), discovered in 1978 at Site G and attributed to Australopithecus afarensis. Here, we report hominin tracks unearthed in the new Site S at Laetoli and referred to two bipedal individuals (S1 and S2) moving on the same palaeosurface and in the same direction as the three hominins documented at Site G. The stature estimates for S1 greatly exceed those previously reconstructed for Au. afarensis from both skeletal material and footprint data. In combination with a comparative reappraisal of the Site G footprints, the evidence collected here embodies very important additions to the Pliocene record of hominin behaviour and morphology. Our results are consistent with considerable body size variation and, probably, degree of sexual dimorphism within a single species of bipedal hominins as early as 3.66 million years ago.

The Acheulian and Early Middle Paleolithic in Latium (Italy): Stability and Innovation.

We present here the results of a technological and typological analysis of the Acheulian and early Middle Paleolithic assemblages from Torre in Pietra (Latium, Italy) together with comparisons with the Acheulian small tools of Castel di Guido. The assemblages were never chronometrically dated before. We have now 40Ar/39Ar dates and ESR-U-series dates, within a geomorphological framework, which support correlations to marine isotope stages. The Acheulian (previously correlated to MIS 9) is now dated to MIS 10 while the Middle Paleolithic is dated to MIS 7. Lithic analyses are preceded by taphonomic evaluations. The Levallois method of the Middle Paleolithic assemblage is an innovation characterized by the production of thin flake blanks without cortex. In contrast, the small tool blanks of the Acheulian were either pebbles or thick flakes with some cortex. They provided a relatively easy manual prehension. The choice of Levallois thin flake blanks in the Middle Paleolithic assemblage suggest that the new technology is most likely related to the emergence of hafting. Accordingly, the oldest direct evidence of hafting technology is from the site of Campitello Quarry in Tuscany (Central Italy) where birch-bark tar, found on the proximal part of two flint flakes, is dated to the end of MIS 7. Nevertheless, a peculiar feature of the Middle Paleolithic at Torre in Pietra is the continuous presence of small tool blanks on pebbles and cores and on thick flake albeit at a much lower frequency than in the older Acheulian industries. The adoption of the new technology is thus characterized by innovation combined with a degree of stability. The persistence of these habits in spite of the introduction of an innovative technique underlies the importance of cultural transmission and conformity in the behavior of Neandertals.

A new chronostratigraphic framework for the Upper Palaeolithic of Riparo Mochi (Italy).

The rockshelter of Mochi, on the Ligurian coast of Italy, is often used as a reference point in the formation of hypotheses concerning the arrival of the Aurigancian in Mediterranean Europe. Yet, the site is poorly known. Here, we describe the stratigraphic sequence based on new field observations and present 15 radiocarbon determinations from the Middle Palaeolithic (late Mousterian) and Early Upper Palaeolithic (Aurignacian and Gravettian) levels. The majority of dates were produced on humanly modified material, specifically marine shell beads, which comprise some of the oldest directly-dated personal ornaments in Europe. The radiocarbon results are incorporated into a Bayesian statistical model to build a new chronological framework for this key Palaeolithic site. A tentative correlation of the stratigraphy to palaeoclimatic records is also attempted.

Paleoenvironmental analysis.

New analysis has been carried out concerning the palaeoenvironmental reconstruction of some Italian sites dating from the Middle Pleistocene to the Bronze Age. Different aspects have been investigated on each site considering the data collected. The following sites have been analyzed: Isernia La Pineta (Molise); Visogliano and Caverna degli Orsi (Tieste); Toirano Caves (Liguria); Grotta Paglicci (Gargano); Riparo del Molare (Salerno); Grotta del Cavallo (Lecce); Castellaro Lagusello (Monzambano, Mantova).

Inorganic raw materials economy and provenance of chipped industry in some stone age sites of northern and central Italy.

An opportunistic and local choice of raw materials is typically attested in the Lower and Middle Paleolithic industries throughout Italy. The quality of the raw material usually affected the flaking technology and quality of the products. In the Upper Paleolithic and the Mesolithic, raw material procurement strategies were more complex. Flint was exploited both locally, in areas where abundant outcrops of raw materials were available (such as the Lessini mountains), and in distant localities, after which it was transported or exchanged over medium/long distances. Different routes of exchange were thus followed in the various periods; good reconstruction of these routes have been provided by a study of the Garfagnana sites in Northern Tuscany, and the Mesolithic deposit of Mondeval de Sora (Dolomites). An interesting example of a Late Upper Paleolithic flint quarry and workshop were found in Abruzzo, in the San Bartolomeo shelter. The extended trade of obsidian from Lipari, Palmarola and Sardinia to the Italian Peninsula is attested in the Neolithic, with some differences concerning the age and different areas.

Living-floors and structures from the Lower Paleolithic to the Bronze Age in Italy.

New researches have been performed on the analysis of some Italian dwelling structures dating from the Lower Paleolithic to Bronze Age. Different methods have been applied to each study according to the extensions of the areas explored. The following sites have been analyzed: Isernia La Pineta (Molise), Visogliano (Trieste) - Lower Paleolithic; Grotta del Cavallo (Lecce), Grotta Grande and Riparo del Molare (Salerno) - Middle Paleolithic; Grotta di Fumane (Verona), Riparo Tagliente (Verona), Grotta Continenza (Fucino L'Aquila), San Bartolomeo (Maiella Mountain, Abruzzo) - Upper Paleolithic; Mondeval de Sora (Belluno), Alpe Veglia (Verbania) and Grotta Edera (Aurisina, Trieste) -Mesolithic; Cala Giovanna Piano (Pianosa Island, Livorno), Contraguda (Perfugas, Sassari), Colle Santo Stefano (Fucino, L'Aquila), Catignano (Pescara), Settefonti (L'Aquila) - Neolithic; Castellaro Lagusello (Monzambano, Mantua) - Bronze Age.