Quantifying hominin morphological diversity at the end of the middle Pleistocene: Implications for the origin of Homo sapiens.

OBJECTIVES: The Middle Pleistocene (MP) saw the emergence of new species of hominins: Homo sapiens in Africa, H. neanderthalensis, and possibly Denisovans in Eurasia, whose most recent common ancestor is thought to have lived in Africa around 600 ka ago. However, hominin remains from this period present a wide range of morphological variation making it difficult to securely determine their taxonomic attribution and their phylogenetic position within the Homo genus. This study proposes to reconsider the phenetic relationships between MP hominin fossils in order to clarify evolutionary trends and contacts between the populations they represent. MATERIALS AND METHODS: We used a Geometric Morphometrics approach to quantify the morphological variation of the calvarium of controversial MP specimens from Africa and Eurasia by using a comparative sample that can be divided into 5 groups: H. ergaster, H. erectus, H. neanderthalensis, and H. sapiens, as well as individuals from current modern human populations. We performed a Generalized Procrustes Analysis, a Principal Component Analysis, and Multinomial Principal Component Logistic Regressions to determine the phenetic affinities of the controversial Middle Pleistocene specimens with the other groups. RESULTS: MP African and Eurasian specimens represent several populations, some of which show strong affinities with H. neanderthalensis in Europe or H. sapiens in Africa, others presenting multiple affinities. DISCUSSION: These MP populations might have contributed to the emergence of these two species in different proportions. This study proposes a new framework for the human evolutionary history during the MP.

What do brain endocasts tell us? A comparative analysis of the accuracy of sulcal identification by experts and perspectives in palaeoanthropology.

Palaeoneurology is a complex field as the object of study, the brain, does not fossilize. Studies rely therefore on the (brain) endocranial cast (often named endocast), the only available and reliable proxy for brain shape, size and details of surface. However, researchers debate whether or not specific marks found on endocasts correspond reliably to particular sulci and/or gyri of the brain that were imprinted in the braincase. The aim of this study is to measure the accuracy of sulcal identification through an experiment that reproduces the conditions that palaeoneurologists face when working with hominin endocasts. We asked 14 experts to manually identify well-known foldings in a proxy endocast that was obtained from an MRI of an actual in vivo Homo sapiens head. We observe clear differences in the results when comparing the non-corrected labels (the original labels proposed by each expert) with the corrected labels. This result illustrates that trying to reconstruct a sulcus following the very general known shape/position in the literature or from a mean specimen may induce a bias when looking at an endocast and trying to follow the marks observed there. We also observe that the identification of sulci appears to be better in the lower part of the endocast compared to the upper part. The results concerning specific anatomical traits have implications for highly debated topics in palaeoanthropology. Endocranial description of fossil specimens should in the future consider the variation in position and shape of sulci in addition to using models of mean brain shape. Moreover, it is clear from this study that researchers can perceive sulcal imprints with reasonably high accuracy, but their correct identification and labelling remains a challenge, particularly when dealing with extinct species for which we lack direct knowledge of the brain.

Investigating the relationship between cranial bone thickness and diploic channels: A first comparison between fossil Homo sapiens and Homo neanderthalensis.

Diploic veins are part of the circulatory system of the head. They transport venous blood and cerebrospinal fluid and are housed in diploic channels (DCs). DCs are highly variable in terms of their position, extension, and size. These parameters were hypothesized to be related to the variations in cranial vault thickness (CVT). For the first time, we analyzed the spatial relationship between CVT and DCs in a sample of eight H. neanderthalensis and H. sapiens cranial fossils. Using micro-CT scanning data, we constructed color maps of the CVT and visually inspected whether the regional thickness variation was associated with the morphology and distribution of the DC branches. The results showed that when regional bone thickness was below a certain threshold, no DCs or scattered small DC branches were present. Larger DC branches appeared only when the thickness exceeded the threshold. However, once the threshold was reached, further increases in thickness no longer resulted in more or larger DCs. This study also found that our sample of H. neanderthalensis and H. sapiens have different distribution patterns in thin areas, which may affect how their DCs connect with different branches of the middle meningeal vessels. This preliminary study provides evidence for the discussion on the interaction between the cranium, brain, and blood vessels. Future research should include more hominin fossils to better document the variation within each species and possible evolutionary trends among hominin lineages.

The diploic venous system in Homo neanderthalensis and fossil Homo sapiens: A study using high-resolution computed tomography.

OBJECTIVES: The diploic venous system has been hypothesized to be related to human brain evolution, though its evolutionary trajectory and physiological functions remain largely unclear. This study examines the characteristics of the diploic venous channels (DCs) in a selection of well-preserved Homo neanderthalensis and Upper Paleolithic Homo sapiens crania, searching for the differences between the two taxa and exploring the associations between brain anatomy and DCs. MATERIALS AND METHODS: Five H. neanderthalensis and four H. sapiens fossil specimens from Western Europe were analyzed. Based on Micro-CT scanning and 3D reconstruction, the distribution pattern and draining orifices of the DCs were inspected qualitatively. The size of the DCs was quantified by volume calculation, and the degree of complexity was quantified by fractal analyses. RESULTS: High-resolution data show the details of the DC structures not documented in previous studies. H. neanderthalensis and H. sapiens specimens share substantial similarities in the DCs. The noticeable differences between the two samples manifest in the connecting points surrounding the frontal sinuses, parietal foramina, and asterional area. DISCUSSION: This study provides a better understanding of the anatomy of the DCs in H. neanderthalensis and H. sapiens. The connection patterns of the DCs have potential utility in distinguishing between the two taxa and in the phylogenetic and taxonomic discussion of the Neandertal-like specimens with controversial taxonomic status.

Lateralized behaviors in living humans: Application in the context of hominin brain evolution.

The left and right hemispheres of our brains differ subtly in structure, and each is dominant in processing specific cognitive tasks. Our species has a unique system of distributing behavior and cognition between each cerebral hemisphere, with a preponderance of pronounced side biases and lateralized functions. This hemisphere-dependent relationship between cognitive, sensory or motor function and a set of brain structures is called hemispheric specialization. Hemispheric specialization has led to the emergence of model systems to link anatomical asymmetries to brain function and behavior. Scientific research on hemispheric specialization and lateralized functions in living humans focuses on three major domains: (1) hand preferences, (2) language, and (3) visuospatial skills and attention. In this chapter we present an overview of this research with a specific focus on living humans and the applications of this research in the context of hominin brain evolution. Our objective is to put into perspective what we know about brain-behavior relationships in living humans and how we can apply the same methods to investigate this relationship in fossil hominin species, and thus improve our understanding of the emergence and development of complex cognitive abilities.

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.

The Internal Cranial Anatomy of a Female With Endocrine Disorders From a Mediaeval Population.

Gigantism and acromegaly have been observed in past populations; however, analyses usually focus on the morphological features of the post-cranial skeleton. The aim of this study is to characterize the internal anatomical features of the skull (brain endocast anatomy and asymmetry, frontal pneumatization, cranial thickness, sella turcica size) of an adult individual from the 11-14th centuries with these two diseases, in comparison with non-pathological individuals from the same population. The material consisted of 33 adult skulls from a mediaeval population, one of them belonging to an adult female with endocrine disorders (OL-23/77). Based on the CT scans, the internal cranial anatomy was analysed. The sella turcica of OL-23/77 is much larger than in the comparative sample. The endocast of the individual OL-23/77 shows a left frontal/left occipital petalia, while the comparative population mostly had right frontal/left occipital petalias. The asymmetry in petalia location in OL-23/77 comes within the range of variation observed in the comparative population. The individual has high values for cranial thickness. The frontal sinuses of the specimen analysed are similar in size and shape to the comparative sample only for data scaled to the skull length. Enlarged sella turcica is typical for individuals with acromegaly/gigantism. The pattern of the left frontal/left occipital petalia in the specimen OL-23/77 is quite rare. The position of the endocranial petalias has not influenced the degree of asymmetry in the specimen. Despite the large bone thickness values, skull of OL-23/77 does not show any abnormal features. The skull/endocast relationship in this individual shows some peculiarities in relation to its large size, while other internal anatomical features are within the normal range of variation of the comparative sample.

Nasopharyngeal morphology contributes to understanding the "muddle in the middle" of the Pleistocene hominin fossil record.

The late archeologist Glynn Isaac first applied the term "muddle in the middle" to a poorly understood period in the Middle Pleistocene human fossil record. This study uses the nasopharyngeal boundaries as a source of traits that may inform this unclear period of human evolution. The nasopharynx lies at the nexus of several vital physiological systems, yet relatively little is known about its importance in human evolution. We analyzed a geographically diverse contemporary Homo sapiens growth series (n = 180 adults, 237 nonadults), Homo neanderthalensis (La Chapelle aux Saints 1, La Ferrassie 1, Forbes Quarry 1, Monte Circeo 1, and Saccopastore 1), mid-Pleistocene Homo (Atapuerca 5, Kabwe 1, Petralona 1, and Steinheim 1), and two Homo erectus sensu lato (KNM-ER 3733 and Sangiran 17). Methods include traditional (Analysis 1) and 3D geometric morphometric analysis (Analysis 2). H. erectus exhibited tall, narrow nasopharyngeal shape, a robust, ancestral morphology. Kabwe 1 and Petralona 1 plotted among H. sapiens in Analysis 2, exhibiting relatively shorter and vertical cartilaginous Eustachian tubes and vertical medial pterygoid plates. Atapuerca 5 and Steinheim 1 exhibited horizontal vomeral orientation similar to H. neanderthalensis, indicating greater relative soft palate length and anteroposterior nasopharynx expansion. They may exhibit synapomorphies with H. neanderthalensis, supporting the accretionary hypothesis. Species-level differences were found among H. sapiens and H. neanderthalensis, including relatively longer dilator tubae muscles and extreme facial airorhynchy among Neanderthals. Furthermore, H. neanderthalensis were autapomorphic in exhibiting horizontal pterygoid plate orientation similar to human infants, suggesting that they may have had inferiorly low placement of the torus tubarius and Eustachian tube orifice on the lateral nasopharyngeal wall in life. This study supports use of osseous nasopharyngeal boundaries both for morphological characters and understanding evolution of otitis media susceptibility in living humans.

Browsing Multiple Subjects When the Atlas Adaptation Cannot Be Achieved via a Warping Strategy.

Brain mapping studies often need to identify brain structures or functional circuits into a set of individual brains. To this end, multiple atlases have been published to represent such structures based on different modalities, subject sets, and techniques. The mainstream approach to exploit these atlases consists in spatially deforming each individual data onto a given atlas using dense deformation fields, which supposes the existence of a continuous mapping between atlases and individuals. However, this continuity is not always verified, and this "iconic" approach has limits. We present in this study an alternative, complementary, "structural" approach, which consists in extracting structures from the individual data, and comparing them without deformation. A "structural atlas" is thus a collection of annotated individual data with a common structure nomenclature. It may be used to characterize structure shape variability across individuals or species, or to train machine learning systems. This study exhibits Anatomist, a powerful structural 3D visualization software dedicated to building, exploring, and editing structural atlases involving a large number of subjects. It has been developed primarily to decipher the cortical folding variability; cortical sulci vary enormously in both size and shape, and some may be missing or have various topologies, which makes iconic approaches inefficient to study them. We, therefore, had to build structural atlases for cortical sulci, and use them to train sulci identification algorithms. Anatomist can display multiple subject data in multiple views, supports all kinds of neuroimaging data, including compound structural object graphs, handles arbitrary coordinate transformation chains between data, and has multiple display features. It is designed as a programming library in both C++ and Python languages, and may be extended or used to build dedicated custom applications. Its generic design makes all the display and structural aspects used to explore the variability of the cortical folding pattern work in other applications, for instance, to browse axonal fiber bundles, deep nuclei, functional activations, or other kinds of cortical parcellations. Multimodal, multi-individual, or inter-species display is supported, and adaptations to large scale screen walls have been developed. These very original features make it a unique viewer for structural atlas browsing.

Variation in cross-sectional indicator of femoral robusticity in Homo sapiens and Neandertals.

Variations in the cross-sectional properties of long bones are used to reconstruct the activity of human groups and differences in their respective habitual behaviors. Knowledge of what factors influence bone structure in Homo sapiens and Neandertals is still insufficient thus, this study investigated which biological and environmental variables influence variations in the femoral robusticity indicator of these two species. The sample consisted of 13 adult Neandertals from the Middle Paleolithic and 1959 adult individuals of H. sapiens ranging chronologically from the Upper Paleolithic to recent times. The femoral biomechanical properties were derived from the European data set, the subject literature, and new CT scans. The material was tested using a Mantel test and statistical models. In the models, the polar moment of area (J) was the dependent variable; sex, age, chronological period, type of lifestyle, percentage of the cortical area (%CA), the ratio of second moment areas of inertia about the X and Y axes (Ix/Iy), and maximum slope of the terrain were independent covariates. The Mantel tests revealed spatial autocorrelation of the femoral index in H. sapiens but not in Neandertals. A generalized additive mixed model showed that sex, %CA, Ix/Iy, chronological period, and terrain significantly influenced variation in the robusticity indicator of H. sapiens femora. A linear mixed model revealed that none of the analyzed variables correlated with the femoral robusticity indicator of Neandertals. We did not confirm that the gradual decline in the femoral robusticity indicator of H. sapiens from the Middle Paleolithic to recent times is related to the type of lifestyle; however, it may be associated with lower levels of mechanical loading during adolescence. The lack of correlation between the analysed variables and the indicator of femoral robusticity in Neandertals may suggest that they needed a different level of mechanical stimulus to produce a morphological response in the long bone than H. sapiens.

The cranial base and related internal anatomical features in Homo neanderthalensis and Homo sapiens.

The cranial anatomy of Homo neanderthalensis and Homo sapiens is well documented in the paleoanthropological and medical literature. However, there are few high-quality visual guides of their comparative morphology. We give here a detailed description of the anatomy of two important fossil specimens, La Chapelle-aux-Saints 1 and abri Pataud 1, based on high-resolution imaging data with each specimen representing the respective morphologies of H. neanderthalensis and H. sapiens. We describe the comparative morphology of external, endocranial, and internal characteristics of the cranium, with a focus on the petrous and tympanic portions of the temporal bone. This descriptive approach shows differences between our specimens, including in positions of cerebral components relative to cranial structures and patterns of dural sinus drainage. Numerous external and internal differences in the shape of the petrous temporal are also described, including its articulation with the tympanic bone and the orientation of the petrotympanic crest. The presence of a large protuberance between the osseous Eustachian tube orifice and carotid foramen in H. neanderthalensis suggests that the levator veli palatini muscle took origin more laterally than the dilator tubae arm of the tensor veli palatini muscle, a feature shared with H. sapiens. The overall pattern that emerges is one in which two species have undergone large-scale evolutionary changes in a functionally critical region. Such differences necessitate high-quality visualization and consideration of both internal and external morphology.

Covariation of proximal finger and toe phalanges in Homo sapiens: A novel approach to assess covariation of serially corresponding structures.

OBJECTIVES: As hands and feet are serially repeated corresponding structures in tetrapods, the morphology of fingers and toes is expected to covary due to a shared developmental origin. The present study focuses on the covariation of the shape of proximal finger and toe phalanges of adult Homo sapiens to determine whether covariation is different in the first ray relative to the others, as its morphology is also different. MATERIAL AND METHODS: Proximal phalanges of 76 individuals of unknown sex (Muséum national d'Histoire naturelle, Paris, and the Natural History Museum, London) were digitized using a surface scanner. Landmarks were positioned on 3D surface models of the phalanges. Generalized Procrustes analysis and two-block partial least squares (PLS) analyses were conducted. A novel landmark-based geometric morphometric approach focusing on covariation is based on a PCoA of the angles between PLS axes in morphospace. The results can be statistically evaluated. RESULTS: The difference in PCo scores between the first and the other rays indicates that the integration between the thumb and the big toe is different from that between the lateral rays of the hand and foot. DISCUSSION: We speculate that the results are possibly the evolutionary consequence of differential selection pressure on the big toe relative to the other toes related to the rise of bipedalism, which is proposed to have emerged very early in the hominin clade. In contrast, thumb morphology and its precision grip never ceased undergoing changes, suggesting less acute selection pressures related to the evolution of the precision grip.

The endocast of the late Middle Paleolithic Manot 1 specimen, Western Galilee, Israel.

Studying endocasts has long allowed anthropologists to examine changes in the external topography and the overall size of the brain throughout the evolutionary history of hominins. The nearly complete calvaria of Manot 1 presents an opportunity to gain insights into the external brain morphology, vascular system, and dimensions of the brain of this late Middle Paleolithic hominin. Detailed size and shape analyses of the Manot 1 endocast indicate a modern Homo sapiens anatomy, despite the presence of some primitive features of the calvaria. Traits considered to be derived endocranial features for H. sapiens are present in Manot 1, including an elongated parietal sagittal chord with an elevated superior part of the hemisphere, a widened posterior part of the frontal lobes, a considerable development of the parietal reliefs such as the supramarginal lobules, and a slight posterior projection of the occipital lobes. These findings, together with data presented in previous studies, rule out the possibility of a direct Neanderthal ancestry for the Manot 1 hominin and instead confirm its affiliation with H. sapiens. The Manot 1 calvaria is more similar to that of later Upper Paleolithic H. sapiens than it is to the earlier Levantine populations of Skhul and Qafzeh. The late Middle Paleolithic date of Manot 1 provides an opportunity to analyze the recent developments in human cerebral morphology and organization.

Pluridisciplinary evidence for burial for the La Ferrassie 8 Neandertal child.

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

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.

The human remains from Axlor (Dima, Biscay, northern Iberian Peninsula).

OBJECTIVES: We provide the description and comparative analysis of all the human fossil remains found at Axlor during the excavations carried out by J. M. de Barandiarán from 1967 to 1974: a cranial vault fragment and seven teeth, five of which likely belonged to the same individual, although two are currently lost. Our goal is to describe in detail all these human remains and discuss both their taxonomic attribution and their stratigraphic context. MATERIALS AND METHODS: We describe external and internal anatomy, and use classic and geometric morphometrics. The teeth from Axlor are compared to Neandertals, Upper Paleolithic, and recent modern humans. RESULTS: Two teeth (a left dm2 , a left di1 ) and the parietal fragment show morphological features consistent with a Neandertal classification, and were found in an undisturbed Mousterian context. The remaining three teeth (plus the two lost ones), initially classified as Neandertals, show morphological features and a general size that are more compatible with their classification as modern humans. DISCUSSION: A left parietal fragment (Level VIII) from a single probably adult Neandertal individual was recovered during the old excavations performed by Barandiarán. Additionally, two different Neandertal children lost deciduous teeth during the formations of levels V (left di1 ) and IV (right dm2 ). In addition, a modern human individual is represented by five remains (two currently lost) from a complex stratigraphic setting. Some of the morphological features of these remains suggest that they may represent one of the scarce examples of Upper Paleolithic modern human remains in the northern Iberian Peninsula, which should be confirmed by direct dating.

Neandertal-like traits visible in the internal structure of non-supranuchal fossae of some recent Homo sapiens: The problem of their identification in hominins and phylogenetic implications.

Although recently the internal structure of the non-supranuchal fossa of Homo sapiens has been described and compared to that observed in the Neandertal suprainiac fossa, until now it has not been examined in any modern human children. In this study, the internal structure of this fossa in the occipital bones of three children (two aged 3‒4 years and one aged 5 years ± 16 months) and one adult individual representing recent Homo sapiens from Australia was analysed and compared to that of the Neandertal suprainiac fossa. In order to analyse the internal composition of the fossae of the examined specimens, initially, high-resolution micro-CT datasets were obtained for their occipital bones; next, 3D topographic maps of the variation in thickness of structural layers of the occipital bones were made and 2D virtual sections in the median region of these fossae were prepared. In the fossa of one immature individual, the thinning of the diploic layer characteristic of a Neandertal suprainiac fossa was firmly diagnosed. The other Neandertal-like trait, concerning the lack of substantial thinning of the external table of the bone in the region of the fossa, was established in two individuals (one child and one adult) due to the observation of an irregular pattern of the thickness of this table in the other specimens, suggesting the presence of an inflammatory process. Our study presents, for the first time, Neandertal-like traits (but not the whole set of features that justifies the autapomorphic status of the Neandertal supraniac fossa) in the internal structure of non-supranuchal fossae of some recent Homo sapiens. We discuss the phylogenetic implications of the results of our analysis and stress the reasons that use of the 3D topographic mapping method is important for the correct diagnosis of Neandertal traits of the internal structure of occipital fossae.

Isolated teeth from La Ferrassie: Reassessment of the old collections, new remains, and their implications.

OBJECTIVES: We provide the description and comparative analysis of six new teeth from the site of La Ferrassie. Our goal is to discuss their taxonomic attribution, and to provide an updated inventory of Neandertal and modern human remains from La Ferrassie in their associated archeological context. MATERIALS AND METHODS: We use external and internal anatomy, classic morphometrics, and geometric morphometrics. The teeth from La Ferrassie are compared to several samples of contemporary Neandertals and upper Paleolithic modern humans and to recent modern humans. RESULTS: Three specimens are classified as Neandertals, two as modern humans, and one remains unclassified. DISCUSSION: Based on the previously known fossil samples and the new teeth reported here, there are currently a minimum of four adult and five immature Neandertal individuals coming from the "Grand Abri" and a minimum of two modern human adult individuals: one from "Grand Abri" and one from "Grotte." It is noteworthy that the spatial distribution of the recovered Neandertal remains is not restricted to the area where the LF1-LF 8 were found but now covers the full extension of the excavated area. Moreover, while both Neandertal and modern human occupations have yielded isolated human remains, the partial-to-complete skeletons only belong to Neandertals. These considerations open new perspectives for the understanding of the occupation and use of the La Ferrassie site.