Reassessment of the human mandible from Banyoles (Girona, Spain).

Since the discovery of a human mandible in 1887 near the present-day city of Banyoles, northeastern Spain, researchers have generally emphasized its archaic features, including the lack of chin structures, and suggested affinities with the Neandertals or European Middle Pleistocene (Chibanian) specimens. Uranium-series and electron spin resonance dating suggest the mandible dates to the Late Pleistocene (Tarantian), approximately ca. 45-66 ka. In this study, we reassessed the taxonomic affinities of the Banyoles mandible by comparing it to samples of Middle Pleistocene fossils from Africa and Europe, Neandertals, Early and Upper Paleolithic modern humans, and recent modern humans. We evaluated the frequencies and expressions of morphological features and performed a three-dimensional geometric morphometric analysis on a virtual reconstruction of Banyoles to capture overall mandibular shape. Our results revealed no derived Neandertal morphological features in Banyoles. While a principal component analysis based on Euclidean distances from the first two principal components clearly grouped Banyoles with both fossil and recent Homo sapiens individuals, an analysis of the Procrustes residuals demonstrated that Banyoles did not fit into any of the comparative groups. The lack of Neandertal features in Banyoles is surprising considering its Late Pleistocene age. A consideration of the Middle Pleistocene fossil record in Europe and southwest Asia suggests that Banyoles is unlikely to represent a late-surviving Middle Pleistocene population. The lack of chin structures also complicates an assignment to H. sapiens, although early fossil H. sapiens do show somewhat variable development of the chin structures. Thus, Banyoles represents a non-Neandertal Late Pleistocene European individual and highlights the continuing signal of diversity in the hominin fossil record. The present situation makes Banyoles a prime candidate for ancient DNA or proteomic analyses, which may shed additional light on its taxonomic affinities.

The dental remains from the Early Upper Paleolithic of Manot Cave, Israel.

This study presents the dental remains discovered at Manot Cave (MC), Western Galilee, Israel. The cave contains evidence for human occupation during the Early Upper Paleolithic period (46-33 ka) mainly of Early Ahmarian (∼46-42 ka) and Levantine Aurignacian (∼38-34 ka) cultural levels. Six teeth (three deciduous and three permanent) were found at the site, of which four could be thoroughly analyzed. The morphology of the teeth was qualitatively described and analyzed using traditional and geometric morphometric methods. A large comparative sample was used in order to assess the morphological affiliation of the Manot specimens with other Homo groups. The results provided equivocal signals: the upper first premolar (MC-9 P3) is probably modern human; the upper deciduous second molar (MC-10 dm2) and the upper second permanent molar (MC-8 M2) might be modern humans; the lower second deciduous molar (MC-7 dm2) might be Neanderthal. Owing to the small sample size and the almost total lack of distinctive characteristics, our outcome could not supply conclusive evidence to address the question of whether Manot Aurignacian population came from Europe or descended from the local Ahmarian population.

Fossil hominin radii from the Sima de los Huesos Middle Pleistocene site (Sierra de Atapuerca, Spain).

Complete radii in the fossil record preceding recent humans and Neandertals are very scarce. Here we introduce the radial remains recovered from the Sima de los Huesos (SH) site in the Sierra de Atapuerca between 1976 and 2011 and which have been dated in excess of 430 ky (thousands of years) ago. The sample comprises 89 specimens, 49 of which are attributed to adults representing a minimum of seven individuals. All elements are described anatomically and metrically, and compared with other fossil hominins and recent humans in order to examine the phylogenetic polarity of certain radial features. Radial remains from SH have some traits that differentiate them from those of recent humans and make them more similar to Neandertals, including strongly curved shafts, anteroposterior expanded radial heads and both absolutely and relatively long necks. In contrast, the SH sample differs from Neandertals in showing a high overall gracility as well as a high frequency (80%) of an anteriorly oriented radial tuberosity. Thus, like the cranial and dental remains from the SH site, characteristic Neandertal radial morphology is not present fully in the SH radii. We also analyzed the cross-sectional properties of the SH radial sample at two different levels: mid-shaft and at the midpoint of the neck length. When standardized by shaft length, no difference in the mid-shaft cross-sectional properties were found between the SH hominins, Neandertals and recent humans. Nevertheless, due to their long neck length, the SH hominins show a higher lever efficiency than either Neandertals or recent humans. Functionally, the SH radial morphology is consistent with more efficient pronation-supination and flexion-extension movements. The particular trait composition in the SH sample and Neandertals resembles more closely morphology evident in recent human males.

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.

The medial pterygoid tubercle in the Atapuerca Early and Middle Pleistocene mandibles: evolutionary implications.

Numerous studies have attempted to identify the presence of uniquely derived (autoapomorphic) Neandertal features. Here, we deal with the medial pterygoid tubercle (MTP), which is usually present on the internal face of the ascending ramus of Neandertal specimens. Our study stems from the identification of a hypertrophied tubercle in ATD6-96, an Early Pleistocene mandible recovered from the TD6 level of the Atapuerca-Gran Dolina site and attributed to Homo antecessor. Our review of the literature and study of numerous original fossil specimens and high quality replicas confirm that the MTP occurs at a high frequency in Neandertals (ca. 89%) and is also present in over half (ca. 55%) of the Middle Pleistocene Sima de los Huesos (SH) hominins. In contrast, it is generally absent or minimally developed in other extinct hominins, but can be found in variable frequencies (

The Neandertal nature of the Atapuerca Sima de los Huesos mandibles.

The recovery of additional mandibular fossils from the Atapuerca Sima de los Huesos (SH) site provides new insights into the evolutionary significance of this sample. In particular, morphological descriptions of the new adult specimens are provided, along with standardized metric data and phylogenetically relevant morphological features for the expanded adult sample. The new and more complete specimens extend the known range of variation in the Atapuerca (SH) mandibles in some metric and morphological details. In other aspects, the addition of new specimens has made it possible to confirm previous observations based on more limited evidence. Pairwise comparisons of individual metric variables revealed the only significant difference between the Atapuerca (SH) hominins and Neandertals was a more vertical symphysis in the latter. Similarly, principal components analysis of size-adjusted variables showed a strong similarity between the Atapuerca (SH) hominins and Neandertals. Morphologically, the Atapuerca (SH) mandibles show nearly the full complement of Neandertal-derived features. Nevertheless, the Neandertals differ from the Atapuerca (SH) mandibles in showing a high frequency of the H/O mandibular foramen, a truncated, thinned and inverted gonial margin, a high placement of the mylohyoid line at the level of the M3, a more vertical symphysis and somewhat more pronounced expression of the chin structures. Size-related morphological variation in the SH hominins includes larger retromolar spaces, more posterior placement of the lateral corpus structures, and stronger markings associated with the muscles of mastication in larger specimens. However, phylogenetically relevant features in the SH sample are fairly stable and do not vary with the overall size of the mandible. Direct comparison of the enlarged mandibular sample from Atapuerca (SH) with the Mauer mandible, the type specimen of H. heidelbergensis, reveals important differences from the SH hominins, and there is no morphological counterpart of Mauer within the SH sample, suggesting the SH fossils should not be assigned to this taxon. The Atapuerca (SH) mandibles show a greater number of derived Neandertal features, particularly those related to midfacial prognathism and in the configuration of the superior ramus, than other European middle Pleistocene specimens. This suggests that more than one evolutionary lineage co-existed in the middle Pleistocene, and, broadly speaking, it appears possible to separate the European middle Pleistocene mandibular remains into two distinct groupings. One group shows a suite of derived Neandertal features and includes specimens from the sites of Atapuerca (SH), Payre, l'Aubesier and Ehringsdorf. The other group includes specimens that generally lack derived Neandertal features and includes the mandibles from the sites of Mauer, Mala Balanica, Montmaurin and (probably) Visogliano. The two published Arago mandibles differ strongly from one another, with Arago 2 probably belonging to this former group, and Neandertal affinities being more difficult to identify in Arago 13. Outside of the SH sample, derived Neandertal features in the mandible only become more common during the second half of the middle Pleistocene. Acceptance of a cladogenetic pattern of evolution during the European middle Pleistocene has the potential to reconcile the predictions of the accretion model and the two phases model for the appearance of Neandertal morphology. The precise taxonomic classification of the SH hominins must contemplate features from the dentition, cranium, mandible and postcranial skeleton, all of which are preserved at the SH site. Nevertheless, the origin of the Neandertal clade may be tied to a speciation event reflected in the appearance of a suite of derived Neandertal features in the face, dentition and mandible, all of which are present in the Atapuerca (SH) hominins. This same suite of features also provides a useful anatomical basis to include other European middle Pleistocene mandibles and crania within the Neandertal clade.

A symbolic Neanderthal accumulation of large herbivore crania.

This work examines the possible behaviour of Neanderthal groups at the Cueva Des-Cubierta (central Spain) via the analysis of the latter's archaeological assemblage. Alongside evidence of Mousterian lithic industry, Level 3 of the cave infill was found to contain an assemblage of mammalian bone remains dominated by the crania of large ungulates, some associated with small hearths. The scarcity of post-cranial elements, teeth, mandibles and maxillae, along with evidence of anthropogenic modification of the crania (cut and percussion marks), indicates that the carcasses of the corresponding animals were initially processed outside the cave, and the crania were later brought inside. A second round of processing then took place, possibly related to the removal of the brain. The continued presence of crania throughout Level 3 indicates that this behaviour was recurrent during this level's formation. This behaviour seems to have no subsistence-related purpose but to be more symbolic in its intent.

Fossil human remains from Bolomor Cave (Valencia, Spain).

Systematic excavations carried out since 1989 at Bolomor Cave have led to the recovery of four Pleistocene human fossil remains, consisting of a fibular fragment, two isolated teeth, and a nearly complete adult parietal bone. All of these specimens date to the late Middle and early Late Pleistocene (MIS 7-5e). The fibular fragment shows thick cortical bone, an archaic feature found in non-modern (i.e. non-Homo sapiens) members of the genus Homo. Among the dental remains, the lack of a midtrigonid crest in the M(1) represents a departure from the morphology reported for the majority of Neandertal specimens, while the large dimensions and pronounced shoveling of the marginal ridges in the C(1) are similar to other European Middle and late Pleistocene fossils. The parietal bone is very thick, with dimensions that generally fall above Neandertal fossils and resemble more closely the Middle Pleistocene Atapuerca (SH) adult specimens. Based on the presence of archaic features, all the fossils from Bolomor are attributed to the Neandertal evolutionary lineage.

Middle Pleistocene dental remains from Qesem Cave (Israel).

This study presents a description and comparative analysis of Middle Pleistocene permanent and deciduous teeth from the site of Qesem Cave (Israel). All of the human fossils are assigned to the Acheulo-Yabrudian Cultural Complex (AYCC) of the late Lower Paleolithic. The Middle Pleistocene age of the Qesem teeth (400-200 ka) places them chronologically earlier than the bulk of fossil hominin specimens previously known from southwest Asia. Three permanent mandibular teeth (C(1) -P(4) ) were found in close proximity in the lower part of the stratigraphic sequence. The small metric dimensions of the crowns indicate a considerable degree of dental reduction although the roots are long and robust. In contrast, three isolated permanent maxillary teeth (I(2) , C(1) , and M(3) ) and two isolated deciduous teeth that were found within the upper part of the sequence are much larger and show some plesiomorphous traits similar to those of the Skhul/Qafzeh specimens. Although none of the Qesem teeth shows a suite of Neanderthal characters, a few traits may suggest some affinities with members of the Neanderthal evolutionary lineage. However, the balance of the evidence suggests a closer similarity with the Skhul/Qafzeh dental material, although many of these resemblances likely represent plesiomorphous features.

Evolution of M1 crown size and cusp proportions in the genus Homo.

Previous research into tooth crown dimensions and cusp proportions has proved to be a useful way to identify taxonomic differences in Pliocene and Pleistocene fossil hominins. The present study has identified changes in both M(1) crown size and cusp proportions within the genus Homo, with M(1) overall crown size reduction apparently occurring in two main stages. The first stage (a reduction of ca. 17%) is associated with the emergence of Homo ergaster and Homo erectus sensu stricto. The second stage (a reduction of ca. 10%) occurs in Homo sapiens, but the reduced modern human M(1) tooth crown size was only attained in Upper Paleolithic times. The absolute sizes of the individual cusps are highly positively correlated with overall crown size and dental reduction produces a reduction in the absolute size of each of the cusps. Most of the individual cusps scale isometrically with crown size, but the paracone shows a negative allometric relationship, indicating that the reduction in paracone size is less than in the other M(1) cusps. Thus, the phylogenetically oldest cusp in the upper molars also seems to be the most stable cusp (at least in the M(1)). The most striking change in M(1) cusp proportions is a change in the relative size of the areas of the paracone and metacone. The combination of a small relative paracone and a large relative metacone generally characterizes specimens attributed to early Homo, and the presence of this character state in Australopithecus and Paranthropus suggests it may represent the primitive condition for the later part of the hominin clade. In contrast, nearly all later Homo taxa, with the exception of Homo antecessor, show the opposite condition (i.e. a relatively large paracone and a relatively small metacone). This change in the relationship between the relative sizes of the paracone and metacone is related to an isometric reduction of the absolute size of the metacone. This metacone reduction occurs in the context of relative stability in the paracone as crown size decreases. Among later Homo taxa, both Homo heidelbergensis and Homo neanderthalensis show a further reduction of the metacone and an enlargement of the hypocone. Fossil and contemporary H. sapiens samples show a trend toward increasing the relative size of the protocone and decreasing the relative size of the hypocone. In Europe, modern human M(1) cusp proportions are essentially reached during the Upper Paleolithic. Although some variation was documented among the fossil taxa, we suggest that the relative size of the M(1) paracone and metacone areas may be useful for differentiating the earliest members of our genus from subsequent Homo species.

The earliest modern humans outside Africa.

To date, the earliest modern human fossils found outside of Africa are dated to around 90,000 to 120,000 years ago at the Levantine sites of Skhul and Qafzeh. A maxilla and associated dentition recently discovered at Misliya Cave, Israel, was dated to 177,000 to 194,000 years ago, suggesting that members of the Homo sapiens clade left Africa earlier than previously thought. This finding changes our view on modern human dispersal and is consistent with recent genetic studies, which have posited the possibility of an earlier dispersal of Homo sapiens around 220,000 years ago. The Misliya maxilla is associated with full-fledged Levallois technology in the Levant, suggesting that the emergence of this technology is linked to the appearance of Homo sapiens in the region, as has been documented in Africa.