Reconstructing Middle and Upper Paleolithic human mobility in Portuguese Estremadura through laser ablation strontium isotope analysis.

Understanding mobility and landscape use is important in reconstructing subsistence behavior, range, and group size, and it may contribute to our understanding of phenomena such as the dynamics of biological and cultural interactions between distinct populations of Upper Pleistocene humans. However, studies using traditional strontium isotope analysis are generally limited to identifying locations of childhood residence or nonlocal individuals and lack the sampling resolution to detect movement over short timescales. Here, using an optimized methodology, we present highly spatially resolved 87Sr/86Sr measurements made by laser ablation multicollector inductively coupled plasma mass spectrometry along the growth axis of the enamel of two marine isotope stage 5b, Middle Paleolithic Neanderthal teeth (Gruta da Oliveira), a Tardiglacial, Late Magdalenian human tooth (Galeria da Cisterna), and associated contemporaneous fauna from the Almonda karst system, Torres Novas, Portugal. Strontium isotope mapping of the region shows extreme variation in 87Sr/86Sr, with values ranging from 0.7080 to 0.7160 over a distance of c. 50 km, allowing short-distance (and arguably short-duration) movement to be detected. We find that the early Middle Paleolithic individuals roamed across a subsistence territory of approximately 600 km2, while the Late Magdalenian individual parsimoniously fits a pattern of limited, probably seasonal movement along the right bank of the 20-km-long Almonda River valley, between mouth and spring, exploiting a smaller territory of approximately 300 km2. We argue that the differences in territory size are due to an increase in population density during the Late Upper Paleolithic.

The symbolic role of the underground world among Middle Paleolithic Neanderthals.

Cueva de Ardales in Málaga, Spain, is one of the richest and best-preserved Paleolithic painted caves of southwestern Europe, containing over a thousand graphic representations. Here, we study the red pigment in panel II.A.3 of "Sala de las Estrellas," dated by U-Th to the Middle Paleolithic, to determine its composition, verify its anthropogenic nature, infer the associated behaviors, and discuss their implications. Using optical microscopy, scanning electron microscopy coupled with energy dispersive X-ray spectroscopy, micro-Raman spectroscopy, and X-ray diffraction, we analyzed a set of samples from the panel and compared them to natural coloring materials collected from the floor and walls of the cave. The conspicuously different texture and composition of the geological samples indicates that the pigments used in the paintings do not come from the outcrops of colorant material known in the cave. We confirm that the paintings are not the result of natural processes and show that the composition of the paint is consistent with the artistic activity being recurrent. Our results strengthen the hypothesis that Neanderthals symbolically used these paintings and the large stalagmitic dome harboring them over an extended time span.

Modeling the role of voyaging in the coastal spread of the Early Neolithic in the West Mediterranean.

The earliest dates for the West Mediterranean Neolithic indicate that it expanded across 2,500 km in about 300 y. Such a fast spread is held to be mainly due to a demic process driven by dispersal along coastal routes. Here, we model the Neolithic spread in the region by focusing on the role of voyaging to understand better the core elements that produced the observed pattern of dates. We also explore the effect of cultural interaction with Mesolithic populations living along the coast. The simulation study shows that (i) sea travel is required to obtain reasonable predictions, with a minimum sea-travel range of 300 km per generation; (ii) leapfrog coastal dispersals yield the best results (quantitatively and qualitatively); and (iii) interaction with Mesolithic people can assist the spread, but long-range voyaging is still needed to explain the archaeological pattern.

New Middle Pleistocene hominin cranium from Gruta da Aroeira (Portugal).

The Middle Pleistocene is a crucial time period for studying human evolution in Europe, because it marks the appearance of both fossil hominins ancestral to the later Neandertals and the Acheulean technology. Nevertheless, European sites containing well-dated human remains associated with an Acheulean toolkit remain scarce. The earliest European hominin crania associated with Acheulean handaxes are at the sites of Arago, Atapuerca Sima de los Huesos (SH), and Swanscombe, dating to 400-500 ka (Marine Isotope Stage 11-12). The Atapuerca (SH) fossils and the Swanscombe cranium belong to the Neandertal clade, whereas the Arago hominins have been attributed to an incipient stage of Neandertal evolution, to Homo heidelbergensis, or to a subspecies of Homo erectus A recently discovered cranium (Aroeira 3) from the Gruta da Aroeira (Almonda karst system, Portugal) dating to 390-436 ka provides important evidence on the earliest European Acheulean-bearing hominins. This cranium is represented by most of the right half of a calvarium (with the exception of the missing occipital bone) and a fragmentary right maxilla preserving part of the nasal floor and two fragmentary molars. The combination of traits in the Aroeira 3 cranium augments the previously documented diversity in the European Middle Pleistocene fossil record.

The bony labyrinth in the Aroeira 3 Middle Pleistocene cranium.

The discovery of a partial cranium at the site of Aroeira (Portugal) dating to 389-436 ka augments the current sample of Middle Pleistocene European crania and makes this specimen penecontemporaneous with the fossils from the geographically close Atapuerca Sima de los Huesos (SH) and Arago sites. A recent study of the cranium documented a unique combination of primitive and derived features. The Aroeira 3 cranium preserves the right temporal bone, including the petrosal portion. Virtual reconstruction of the bony labyrinth from µCT scans provides an opportunity to examine its morphology. A series of standard linear and angular measures of the semicircular canals and cochlea in Aroeira 3 were compared with other fossil hominins and recent humans. Our analysis has revealed the absence of derived Neandertal features in Aroeira 3. In particular, the specimen lacks both the derived canal proportions and the low position of the posterior canal, two of the most diagnostic features of the Neandertal bony labyrinth, and Aroeira 3 is more primitive in these features than the Atapuerca (SH) sample. One potentially derived feature (low shape index of the cochlear basal turn) is shared between Aroeira 3 and the Atapuerca (SH) hominins, but is absent in Neandertals. The results of our study provide new insights into Middle Pleistocene population dynamics close to the origin of the Neandertal clade. In particular, the contrasting inner ear morphology between Aroeira 3 and the Atapuerca (SH) hominins suggests a degree of demographic isolation, despite the close geographic proximity and similar age of these two sites.

Taphonomic inferences about Middle Pleistocene hominins: The human cranium of Gruta da Aroeira (Portugal).

OBJECTIVES: The aim of this work is to describe the taphonomic signatures of the Aroeira 3 cranium, with a specific focus on cranial breakage, comparing the cranium with other Middle and Upper Pleistocene hominin fossils in order to approximate the cause of death and the biological agencies and geologic processes involved in the taphonomic record of this specimen. Aroeira-3 was recovered from Acheulean layer X of Gruta da Aroeira (Portugal), dated to 390-436 ka. MATERIALS AND METHODS: Taphonomic analyses noted surface modifications employing standard methods. The cranial breakage pattern of Aroeira 3 was analyzed to assess the presence/absence of perimortem (fresh bone) and postmortem (dry bone) fractures and the possible causes of perimortem skull bone fractures. RESULTS: Aroeira 3 presents substantial bone loss of the left supraorbital arch and the outer cranial table of the frontal squama. Most of the fractures present features consistent with postmortem injuries. The fracture to the posterior region of the parietal bone, however, displays features more usually present in perimortem bone fractures. No evidence of anthropogenic activity or of carnivore modification has been identified. None of the expected features of interpersonal conflict are observed. Finally, the bone loss in the frontal squama and the supraorbital arch could be attributed to different agencies, and a traumatic event cannot be totally ruled out as origin of the bone alteration. DISCUSSION: Cannibalism, secondary treatment of the corpse and accumulation induced by carnivores can all be discarded, making an accident the most plausible explanation for the cranial fracture.

A Common Genetic Origin for Early Farmers from Mediterranean Cardial and Central European LBK Cultures.

The spread of farming out of the Balkans and into the rest of Europe followed two distinct routes: An initial expansion represented by the Impressa and Cardial traditions, which followed the Northern Mediterranean coastline; and another expansion represented by the LBK (Linearbandkeramik) tradition, which followed the Danube River into Central Europe. Although genomic data now exist from samples representing the second migration, such data have yet to be successfully generated from the initial Mediterranean migration. To address this, we generated the complete genome of a 7,400-year-old Cardial individual (CB13) from Cova Bonica in Vallirana (Barcelona), as well as partial nuclear data from five others excavated from different sites in Spain and Portugal. CB13 clusters with all previously sequenced early European farmers and modern-day Sardinians. Furthermore, our analyses suggest that both Cardial and LBK peoples derived from a common ancient population located in or around the Balkan Peninsula. The Iberian Cardial genome also carries a discernible hunter-gatherer genetic signature that likely was not acquired by admixture with local Iberian foragers. Our results indicate that retrieving ancient genomes from similarly warm Mediterranean environments such as the Near East is technically feasible.

The Neandertals of northeastern Iberia: new remains from the Cova del Gegant (Sitges, Barcelona).

The present study describes a new juvenile hominin mandible and teeth and a new juvenile humerus from level V of the GP2 gallery of Cova del Gegant (Spain). The mandible (Gegant-5) preserves a portion of the right mandibular corpus from the M1 distally to the socket for the dc mesially, and the age at death is estimated as 4.5-5.0 years. Gegant-5 shows a single mental foramen located under the dm1/dm2 interdental septum, a relatively posterior placement compared with recent hominins of a similar developmental age. The mental foramen in Gegant-5 is also placed within the lower half of the mandibular corpus, as in the previously described late adolescent/adult mandible (Gegant-1) from this same Middle Paleolithic site. The Gegant-5 canine shows pronounced marginal ridges, a distal accessory ridge, and a pronounced distolingual tubercle. The P3 shows a lingually-displaced protoconid cusp tip and a distal accessory ridge. The P4 shows a slightly asymmetrical crown outline, a continuous transverse crest, a mesially placed metaconid cusp tip, a slight distal accessory ridge, and an accessory lingual cusp. The M1 shows a Y5 pattern of cusp contact and a well-developed and deep anterior fovea bounded posteriorly by a continuous midtrigonid crest. Gegant-4 is the distal portion of a left humerus from a juvenile estimated to be between 5 and 7 years old at death. The specimen shows thick cortical bone. Although fragmentary, the constellation of morphological and metric features indicates Neandertal affinities for these specimens. Their spatial proximity at the site and similar ages at death suggest these remains may represent a single individual. The addition of these new specimens brings the total number of Neandertal remains from the Cova del Gegant to five, and this site documents the clearest evidence for Neandertal fossils associated with Middle Paleolithic stone tools in this region of the Iberian Peninsula.

Human-climate interaction during the Early Upper Paleolithic: testing the hypothesis of an adaptive shift between the Proto-Aurignacian and the Early Aurignacian.

The Aurignacian technocomplex comprises a succession of culturally distinct phases. Between its first two subdivisions, the Proto-Aurignacian and the Early Aurignacian, we see a shift from single to separate reduction sequences for blade and bladelet production, the appearance of split-based antler points, and a number of other changes in stone tool typology and technology as well as in symbolic material culture. Bayesian modeling of available (14)C determinations, conducted within the framework of this study, indicates that these material culture changes are coincident with abrupt and marked climatic changes. The Proto-Aurignacian occurs during an interval (ca. 41.5-39.9 k cal BP) of relative climatic amelioration, Greenland Interstadials (GI) 10 and 9, punctuated by a short cold stadial. The Early Aurignacian (ca. 39.8-37.9 k cal BP) predominantly falls within the climatic phase known as Heinrich Stadial (HS) 4, and its end overlaps with the beginning of GI 8, the former being predominantly characterized by cold and dry conditions across the European continent. We use eco-cultural niche modeling to quantitatively evaluate whether these shifts in material culture are correlated with environmental variability and, if so, whether the ecological niches exploited by human populations shifted accordingly. We employ genetic algorithm (GARP) and maximum entropy (Maxent) techniques to estimate the ecological niches exploited by humans (i.e., eco-cultural niches) during these two phases of the Aurignacian. Partial receiver operating characteristic analyses are used to evaluate niche variability between the two phases. Results indicate that the changes in material culture between the Proto-Aurignacian and the Early Aurignacian are associated with an expansion of the ecological niche. These shifts in both the eco-cultural niche and material culture are interpreted to represent an adaptive response to the relative deterioration of environmental conditions at the onset of HS4.

Dental maturational sequence and dental tissue proportions in the early Upper Paleolithic child from Abrigo do Lagar Velho, Portugal.

Neandertals differ from recent and terminal Pleistocene human populations in their patterns of dental development, endostructural (internal structure) organization, and relative tissue proportions. Although significant changes in craniofacial and postcranial morphology have been found between the Middle Paleolithic and earlier Upper Paleolithic modern humans of western Eurasia and the terminal Pleistocene and Holocene inhabitants of the same region, most studies of dental maturation and structural morphology have compared Neandertals only to later Holocene humans. To assess whether earlier modern humans contrasted with later modern populations and possibly approached the Neandertal pattern, we used high-resolution microtomography to analyze the remarkably complete mixed dentition of the early Upper Paleolithic (Gravettian) child from Abrigo do Lagar Velho, Portugal, and compared it to a Neandertal sample, the late Upper Paleolithic (Magdalenian) child of La Madeleine, and a worldwide extant human sample. Some aspects of the dental maturational pattern and tooth endostructural organization of Lagar Velho 1 are absent from extant populations and the Magdalenian specimen and are currently documented only among Neandertals. Therefore, a simple Neandertal versus modern human dichotomy is inadequate to accommodate the morphostructural and developmental variation represented by Middle Paleolithic and earlier Upper Paleolithic populations. These data reinforce the complex nature of Neandertal-modern human similarities and differences, and document ongoing human evolution after the global establishment of modern human morphology.

Symbolic use of marine shells and mineral pigments by Iberian Neandertals.

Two sites of the Neandertal-associated Middle Paleolithic of Iberia, dated to as early as approximately 50,000 years ago, yielded perforated and pigment-stained marine shells. At Cueva de los Aviones, three umbo-perforated valves of Acanthocardia and Glycymeris were found alongside lumps of yellow and red colorants, and residues preserved inside a Spondylus shell consist of a red lepidocrocite base mixed with ground, dark red-to-black fragments of hematite and pyrite. A perforated Pecten shell, painted on its external, white side with an orange mix of goethite and hematite, was abandoned after breakage at Cueva Antón, 60 km inland. Comparable early modern human-associated material from Africa and the Near East is widely accepted as evidence for body ornamentation, implying behavioral modernity. The Iberian finds show that European Neandertals were no different from coeval Africans in this regard, countering genetic/cognitive explanations for the emergence of symbolism and strengthening demographic/social ones.

Formation processes, fire use, and patterns of human occupation across the Middle Palaeolithic (MIS 5a-5b) of Gruta da Oliveira (Almonda karst system, Torres Novas, Portugal).

Gruta da Oliveira features a c. 13 m-thick infilling that includes a c. 6.5 m-thick archaeological deposit (the "Middle Palaeolithic sequence" complex), which Bayesian modelling of available dating results places in MIS 5a (layers 7-14) and MIS 5b (layers 15-25), c. 71,000-93,000 years ago. The accumulation primarily consists of sediment washed in from the slope through gravitational processes and surface dynamics. The coarse fraction derives from weathering of the cave's limestone bedrock. Tectonic activity and structural instability caused the erosional retreat of the scarp face, explaining the large, roof-collapsed rock masses found through the stratification. The changes in deposition and diagenesis observed across the archaeological sequence are minor and primarily controlled by local factors and the impact of humans and other biological agents. Pulses of stadial accumulation-reflected in the composition of the assemblages of hunted ungulates, mostly open-country and rocky terrain taxa (rhino, horse, ibex)-alternate with interstadial hiatuses-during which carbonate crusts and flowstone formed. Humans were active at the cave throughout, but occupation was intermittent, which allowed for limited usage by carnivores when people visited less frequently. During the accumulation of layers 15-25 (c. 85,000-93,000 years ago), the carnivore guild was dominated by wolf and lion, while brown bear and lynx predominate in layers 7-14 (c. 71,000-78,000 years ago). In the excavated areas, conditions for residential use were optimal during the accumulation of layers 20-22 (c. 90,000-92,000 years ago) and 14 (c. 76,000-78,000 years ago), which yielded dense, hearth-focused scatters of stone tools and burnt bones. The latter are ubiquitous, adding to the growing body of evidence that Middle Palaeolithic Neandertals used fire in regular, consistent manner. The patterns of site usage revealed at Gruta da Oliveira are no different from those observed 50,000 years later in comparable early Upper Palaeolithic and Solutrean cave sites of central Portugal.

Middle Paleolithic human remains from the Gruta Da Oliveira (Torres Novas), Portugal.

Additional Middle Paleolithic human remains from layers 17, 18, and 22 of the Gruta da Oliveira, Portugal consist of a proximal manual phalanx 2 (Oliveira 5), a partial postcanine tooth (Oliveira 6), a humeral diaphysis (Oliveira 7), a distal mandibular molar (Oliveira 8), and a mandibular premolar (P(3) ) (Oliveira 9). Oliveira 5, 6, and 8 are unremarkable for Late Pleistocene humans. The Oliveira 7 right humerus is moderately robust or the individual had the stocky body proportions of other European (including Iberian) Neandertals. The Oliveira 9 P(3) has a large and symmetrical crown and lacks a distal accessory ridge and accessory lingual cusps, overlapping both Neandertal and recent human ranges of variation. It contrasts with at least recent human P(3) s in having relatively thin enamel. These join the Oliveira 1 to 4 remains in further documenting early MIS 3 Neandertal morphology in western Iberia.

Organization of residential space, site function variability, and seasonality of activities among MIS 5 Iberian Neandertals.

Whether ethnoarcheological models of hunter-gatherer mobility, landscape use, and structuration of the inhabited space are relevant to the archeology of Neandertals and the Middle Paleolithic remains controversial. The thin lenses of hearth-associated stone tools and faunal remains excavated in sub-complex AS5 of Cueva Antón (Murcia, Spain) significantly advance these debates. Dated to 77.8-85.1 ka, these living floors are interstratified in river-accumulated sands and were buried shortly after abandonment by low-energy inundation events, with minimal disturbance and negligible palimpsest formation. Stone tools were made and ergonomically modified to fit tasks; their spatial distributions and use-wear reveal hearth-focused activities and a division of the inhabited space into resting and working areas. Site function varied with season of the year: units III-i/j1 and III-i/j2-3 record winter visits focused on filleting and hide processing, while woodworking predominated in unit III-b/d, which subsumes visits to the site over the course of at least one winter, one spring, and one summer. These snapshots of Neandertal behavior match expectations derived from the ethnographic and Upper Paleolithic records for the lifeways of hunter-gatherers inhabiting temperate regions with a markedly seasonal climate.

Critical evaluation of in situ analyses for the characterisation of red pigments in rock paintings: A case study from El Castillo, Spain.

Paint technology, namely paint preparation and application procedures, is an important aspect of painting traditions. With the expansion of archaeometric studies and in situ non-destructive analytical methods, a renewal of technological studies is being observed in rock art. In situ analyses have several limitations that are widely discussed in the literature, however. It is not yet clear whether they provide accurate information on paint technology, except under certain conditions. Here, we evaluated digital microscopic and pXRF in situ analyses for the characterisation of a large set of red and yellow paintings from the El Castillo cave, Cantabria, Spain. We have set experiments and used statistical methods to identify differences between paint components and determine factors impacting pXRF measurements. We found that the compositional heterogeneity of the paintings' environment, especially variations in secondary deposits, was responsible for most of the differences observed between the pXRF signals recorded on the paintings. We concluded that the El Castillo cave environment is not suitable for non-destructive technological studies, but that more favourable contexts might exist. Following previous works and our own results, we advocate a combination of both in situ and laboratory invasive analyses for the study of paint composition and paint technology. Our research protocol, based on the comparison of rock paintings, their substrate, experimental paintings and Fe-normalisation of the signals can improve the reliability of pXRF results. We also propose to include more systematic characterisation of rock wall heterogeneity and the use of microscopic analyses in non-destructive approaches.

Frontal sinuses and human evolution.

The frontal sinuses are cavities inside the frontal bone located at the junction between the face and the cranial vault and close to the brain. Despite a long history of study, understanding of their origin and variation through evolution is limited. This work compares most hominin species' holotypes and other key individuals with extant hominids. It provides a unique and valuable perspective of the variation in sinuses position, shape, and dimensions based on a simple and reproducible methodology. We also observed a covariation between the size and shape of the sinuses and the underlying frontal lobes in hominin species from at least the appearance of Homo erectus. Our results additionally undermine hypotheses stating that hominin frontal sinuses were directly affected by biomechanical constraints resulting from either chewing or adaptation to climate. Last, we demonstrate their substantial potential for discussions of the evolutionary relationships between hominin species.

Isotopic evidence for omnivory among European cave bears: Late Pleistocene Ursus spelaeus from the Pestera cu Oase, Romania.

Previous bone collagen carbon and nitrogen isotopic studies of Late Pleistocene European cave bears (Ursus spelaeus) have shown that these bears frequently had low nitrogen isotope values, similar to those of herbivores and indicating either unusual physiology related to hibernation or a herbivorous diet. Isotopic analysis of animal bone from the Pestera cu Oase (Cave with Bones), Romania, shows that most of its cave bears had higher nitrogen isotope values than the associated herbivores and were, therefore, omnivorous. The Oase bears are securely identified as cave bears by both their morphology and DNA sequences. Although many cave bear populations may have behaved like herbivores, the Oase isotopic data demonstrate that cave bears were capable of altering their diets to become omnivores or even carnivores. These data therefore broaden the dietary profile of U. spelaeus and raise questions about the nature of the carnivore guild in Pleistocene Europe.