Les restes humains du Pliocène final et du début du Pléistocène inférieur de Dmanissi, Géorgie (1991-2000). I - Les crânes, D 2280, D 2282, D 2700

Since 1991, several human remains: 5 skulls, 4 mandibles and numerous postcranial fragments have been discovered on the Dmanissi prehistoric open site. It is an exceptional discovery due to the stratigraphical, paleontological and cultural context, which is well known and accurately well dated (Upper Pliocene-Early Pleistocene). Most of the hominids discovered in the level V and VI are dated between 1.81 My (level V) and 1.77 My (level VI) corresponding to a 40,000 years period. The assemblage of fossil human remains is peculiar due to (1) the quality of bone representation (distinct parts of the skeleton are preserved: skull, thorax, upper and lower limbs, belt), (2) the high degree of bone preservation (skulls and long bones are entire, rarely broken or crushed), (3) the diversity age at death estimated for each of the 5 individuals (3 adults, 1 young adult, 1 adolescent of both sexes). The study dealing with the first discovered mandibles and skulls has begun with Leo Gabounia since 1991 and represents several interests: 1) a paleoanthropological interest: the Dmanissi skulls are characterized by their small size; they are short, narrow and low. The skullcaps are less elevated than those of the Homo erectus group and even those of Homo ergaster. They are more elevated than those of Homo habilis and very close to Homo rudolfensis. The elevation and the transversal development of the middle part of the skull in the parietotemporal region are more significant: the Dmanissi specimens are intermediate between Homo habilis and Homo ergaster. In term of cranial capacity, a similar trend is observed. Generally speaking, the skull is slender. The vault is more flat than in Homo erectus, the frontal bone is less developed, divergent and the postorbital constriction is strong. The temporal bone is long, flat and the mastoid part is short. The upper part of the occipital bone is low and narrow. Crests are thin, less developed than in the Homo erectus group. The superior temporal crests are in a high position and a torus angularis is present on the adult-male specimen. The glenoid cavity is large with strong edges. The petrotympanic region is slender with a tympanic circle individualized and it shows a horizontal rotation in a posterior position, which is distinct from Homo erectus. The orthognathic trend of the face distinguishes the Dmanissi specimens from the early Pleistocene hominids (Homo habilis, Homo ergaster) and from the first Eurasian Homo erectus. Nevertheless, the subnasal region of the face is projected. The morphology of the mid-face, showing a developed pillar of the canine, an inframalar incurvation and an anterior position of the root of the zygomaticomaxillary crest, suggests strong masticatory stress. Considering the overall morphology, cranial and metrical features, the Dmanissi fossil skulls are intermediate to the Homo habilis-rudolfensis group and Homo ergaster while they are closer to the former and peculiarly to Homo rudolfensis (ER 1470). However, the Dmanissi fossil skulls are distinct from Homo rudolfensis by numerous features and among them: by their large maximum cranial width (Euryon-Euryon), the posterior rotation of their petrotympanic structure and the strong development of the pillar of their canine. Due to the gracility of their face, the narrowness of their occipital bone, and their cranial base pattern (mastoid region and petrotympanic structure), the Dmanissi fossil skulls are different from the Homo erectus group: 2) the abundance of the human fossils discovered in Dmanissi site provides information about the biodiversity of these hominids with the establishment of the morphological features related to either growth or sexual patterns: 3) compared to modern humans, the Dmanissi fossil skulls seem to follow a different growth pattern. The present study of the fossil skulls discovered is a pioneer step. Indeed, the Dmanissi site has yielded the oldest evidences of the first settlements in Eurasia, which were, until now, attributed to Homo erectus. The Dmanissi fossil skulls are close to the Homo habilis-rudolfensis African group. We attribute these hominids to Homo georgicus.     

A re-examination of the human fossil specimen from Bački Petrovac (Serbia)

A fragmented human calotte was discovered during the early 1950s near Ba?ki Petrovac (Serbia), in association with Palaeolithic stone tools. After its initial publication, the fossil specimen remained largely unknown outside of the Serbian academe and no detailed comparative study has ever been carried out. Since the whereabouts of the fossil itself are currently unknown, and given its potential significance for the Pleistocene human evolution, we re-examine the data published by 0255 and 0260. Using the original measurements, mostly taken on the frontal bone, and a wide comparative sample of 68 fossil specimens, the fossil was compared and analyzed by statistical multivariate methods. We also conducted a visual examination of the morphology based on the available photographic material. Our analysis reveals phenetic similarity with Middle Pleistocene archaic Homo from Africa and anatomically modern Homo sapiens. However, the absence of primitive cranial traits in Ba?ki Petrovac indicates a clear modern Homo sapiens designation. Although lost at the moment, there is a chance for the re-discovery of the fossil in the years to come. This would give us an opportunity to acquire absolute dates and to study the specimen in a more detailed manner.     

Conclusion. Bilan des recherches interdisciplinaires effectuées sur le site de l’Homo erectus de Denizli,Kocabaş, Bassin de Denizli,Anatolie, Turquie. L’Homo erectus aux portes de l’Europe

The study of the site of Kocaba?, which yielded an archaic Homo erectus skullcap, was undertaken in 2011 and 2012, at the request of Professor Mehmet Cihat Alçiçek. This interdisciplinary French–Turkish research programme comprised the geochronological, magnetostratigraphic, biochronological and paleoenvironmental study of the site and the paleoanthropological study of the skullcap itself. The association of large mammals enabled us to attribute the travertine formations bearing the skullcap to the second part of the Upper Pleistocene, and more specifically to between 1.5 and 1.2 million years, because of the disappearance or appearance of certain species. This biochronological age is confirmed by the paleomagnetism study, which places the travertines bearing the skullcap in a period of reversed polarity, underlying a normal polarity formation, which could be attributed to the Cobb Mountain paleomagnetic excursion, dated to 1,194,000 years. The dating of these fauna by the ²⁶Al/¹⁰Be cosmogenic nuclide method by Anne-Elisabeth Lebatard yielded an age older than 1.22 Ma and more recent than 1.5 Ma. The Hominid skullcap from this formation can be attributed to a Homo erectus, slightly more evolved than those of Homo ergaster KNM-ER 3733 (1.78 Ma) and KNM-ER 15,000 (1.5 Ma), similar to that of Daka (Bouri), which is about a million years old and older than the Bodo fossil (estimated at 600,000 years) and Kabwe (between 300,000 and120,000 years). The archaic Homo erectus skullcap from Kocaba?, referred to as Denizli Man, proves that Homo erectus was already present in Anatolia, at the crossroads of Africa, Asia and Europe, a little more than 1.2 million years ago.     

Historique de la découverte et des recherches sur la calotte crânienne d’Homo erectus archaïque de Kocabaş, Bassin de Denizli,Anatolie, Turquie

In November 2002, during the study of the Plio-Pleistocene formations of Denizli Basin, containing significant active and fossil travertine deposits, during a visit to the Dalmersan factory, near the Kocaba? quarries, I identified a Hominid skullcap among bones retrieved during the cutting of travertine slabs. I immediately attributed this skullcap to a Homo erectus. I then handed over the study of this fossil, first to Professor John Kappelman from Texas University, then to a multidisciplinary study led by the Human Paleontology Institute in Paris.     

Mandibular molar root morphology in Neanderthals and Late Pleistocene and recent Homo sapiens

Neanderthals have a distinctive suite of dental features, including large anterior crown and root dimensions and molars with enlarged pulp cavities. Yet, there is little known about variation in molar root morphology in Neanderthals and other recent and fossil members of Homo. Here, we provide the first comprehensive metric analysis of permanent mandibular molar root morphology in Middle and Late Pleistocene Homo neanderthalensis, and Late Pleistocene (Aterian) and recent Homo sapiens. We specifically address the question of whether root form can be used to distinguish between these groups and assess whether any variation in root form can be related to differences in tooth function. We apply a microtomographic imaging approach to visualise and quantify the external and internal dental morphologies of both isolated molars and molars embedded in the mandible (n = 127). Univariate and multivariate analyses reveal both similarities (root length and pulp volume) and differences (occurrence of pyramidal roots and dental tissue volume proportion) in molar root morphology among penecontemporaneous Neanderthals and Aterian H. sapiens. In contrast, the molars of recent H. sapiens are markedly smaller than both Pleistocene H. sapiens and Neanderthals, but share with the former the dentine volume reduction and a smaller root-to-crown volume compared with Neanderthals. Furthermore, we found the first molar to have the largest average root surface area in recent H. sapiens and Neanderthals, although in the latter the difference between M₁ and M₂ is small. In contrast, Aterian H. sapiens root surface areas peak at M₂. Since root surface area is linked to masticatory function, this suggests a distinct occlusal loading regime in Neanderthals compared with both recent and Pleistocene H. sapiens.     

The Neanderthal taxonomic position: models of intra- and inter-specific craniofacial variation

The Neanderthal taxonomic position is a matter of wide disagreement among paleoanthropologists. Some workers consider this fossil human group to represent a different species, Homo neanderthalensis, while others see it as a subspecies of Homo sapiens. This study developed two models of morphological variation to be applied to a comparison between Neanderthals and modern humans: modern human populations provided a measure of intra-specific variation, while the species and subspecies of Pan provided measures of both intra- and inter-specific morphological differences. Although such an approach has been advocated strongly, it has not been systematically undertaken until recently. The techniques of geometric morphometrics were used to collect data in the form of three-dimensional coordinates of craniofacial landmarks. The data were processed using generalized procrustes analysis, and analyzed by an array of multivariate statistical methods, including principal components analysis, canonical variates analysis and Mahalanobis D(2). The morphological distances between Neanderthals and modern humans, and between Neanderthals and Late Paleolithic/early anatomically modern specimens, are consistently greater than the distances among recent human populations, and greater than the distances between the two chimpanzee species. Furthermore, no strong morphological similarities were found between Neanderthals and Late Paleolithic Europeans. This study does not find evidence for Neanderthal contribution to the evolution of modern Europeans. Results are consistent with the recognition of Neanderthals as a distinct species.     

L’homme de Tautavel. Un Homo erectus européen évolué. Homo erectus tautavelensis

One hundred and forty-eight human remains have been discovered during excavations carried out between 1964 and 2014 in la Caune de l’Arago, in Tautavel, in the Eastern Pyrenees. They come from a detailed stratigraphic context comprising 15 archaeostratigraphic units with human presence, with ages ranging from 550 ka for unit Q at the base, to 400 ka for unit C at the top (OIS 14 to 10). During this long period of time, Man experienced two cold and dry climatic periods (stratigraphic complexes I and III), separated by a humid-temperate period (stratigraphic complex II). Most of the human remains come from units F and G, in stratigraphic complex III, associated with a cold and windy steppic environment. The human remains were mixed with the archaeological material and waste from hunted and consumed fauna. The inventory of human remains comprises a majority of cranial elements, and in particular, the anterior portion of a skull, Arago XXI, discovered on 22 July 1971, which revealed the physical aspect of the first Europeans for the first time. The remains are made up of 5 mandibles, 123 teeth (isolated or still on the alveolar arch), several post-cranial skeletal fragments: 9 upper limb elements, 19 lower limb elements, representing 30 deceased individuals, comprising 18 adults and 12 children. The study of these fossils relates them to Asian and African forms of Homo erectus, as they display common characteristics. This observation raises questions as to the existence of this group in Europe. The contribution of the collection of human fossils from Arago thus presents a threefold interest; paleontological, population-based and behavioural. The multiple remains allow us to assess the biodiversity and the composition of this small population. These remains present original features in comparison to Mauer, the classical Homo heidelbergensis European ancestor. The Arago fossils display archaic characteristics not observed on the Mauer mandible; in particular, the great anteroposterior extension of the convex arch toward the fore, the large proportions of the premolars and of the M2, the high robusticity index of the mandible body, the sub-horizontal alveolar planum and the slightly inclined prominent mylohyoidian line. The face of the skull is not yet reduced to allow for the expansion of the brain, a process which develops at a later stage. The skull is low, with an extended frontal, very marked facial prognathism and a powerful masticatory apparatus with temporal crests and a prominent torus angularis, conveying a pantagonal coronal cross-section, contrasting with the regular convexity observed on the skulls from La Sima de los Huesos and Neanderthals. A comparative analysis with the well-documented population discovered in La Sima de los Huesos shows that the latter fossils are more evolved and are more similar to the Neanderthal shape, and at the same time, not very different from the Mauer mandible. The currently known European human fossils point to the following scenario: Homo georgicus, a similar form to the habilis-rudolfensis group, bearing pre-Oldowan and Oldowan technology, was present at the gateways of Europe approximately 1.8 Ma ago. From 1.2–0.8 Ma onwards, the somewhat fragmentary records from Atapuerca, Elefante, Gran Dolina-TD6, could be related to this first lineage. The first Homo erectus with biface cultures to leave Africa arrived at the gateways of Europe some 1.2 Ma ago, as shown by the discovery of the Kocabaş skull cap in Anatolia, similar to the fossils from Buia in Eritrea and Daka in Ethiopia, dated to about 1 Ma. From 0.55 Ma, the set of 148 human remains, and in particular the skull Arago XXI, point to the presence of a new, well-documented form (distinct from Mauer). We propose to relate these remains to Homo erectus tautavelensis, thereby giving this subspecies a geographic connotation. The morphofunctional and cultural characteristics of Homo erectus tautavelensis represent the stock of a long European lineage, leading to the emergence of Neanderthals.     

Datation des travertins de Kocabaş par la méthode des nucléides cosmogéniques Al/Be

The archaic Homo erectus Kocaba? skullcap was discovered at Kocaba?, Denizli, Turkey in travertine formations over- and underlain by conglomerate formations. These units dated by the ²⁶Al/¹⁰Be cosmogenic nuclide method constrained the skullcap age between 1.0 and 1.6 Ma.     

A new brain endocast of Homo erectus from Hulu Cave, Nanjing, China

A new brain endocast of Homo erectus from Hulu Cave, Tangshan, Nanjing is described and compared with a broad sample of endocasts of H. erectus, Neanderthals, and recent modern humans. The Nanjing 1 endocast is reconstructed based on two portions of endocranial casts taken from the original fossil fragments. The fossil was discovered in 1993, near Nanjing, South China and is dated to ～ 0.58-0.62 Ma. The cranial capacity is ～ 876 cc, as determined by endocast water displacement. There are some common features of Nanjing 1 and other H. erectus endocasts that differentiate them from the Neanderthals and modern humans in our sample. These include small cranial capacity, low height dimensions, simple middle meningeal vessel patterns, a high degree of cerebral-over-cerebellar lobe overhang, elongated and quite separated cerebellar lobes, and a narrow, low, short and flat frontal region. Some features are found to vary among H. erectus, Neanderthals and modern humans, such as the lateral Sylvian fissure position and the venous sinus and petalial patterns. The Nanjing 1 endocast has unique, large, superior frontal convolutions, and strongly protruding Broca's caps. In contrast to other Chinese H. erectus from Hexian and Zhoukoudian, Nanjing 1 lacks strong posterior projection of the occipital lobes. Bivariate and principal component analyses indicate that the small volume and shape of Nanjing 1 is most similar to KNM-WT 15000, KNM-ER 3883, Sangiran 2 and Hexian, illustrating the combination of narrow, low, and short frontal lobes with wide posterior lobes.     

A geometric morphometric analysis of hominin upper second and third molars, with particular emphasis on European Pleistocene populations

The study of dental morphology by means of geometric morphometric methods allows for a detailed and quantitative comparison of hominin species that is useful for taxonomic assignment and phylogenetic reconstruction. Upper second and third molars have been studied in a comprehensive sample of Plio- and Pleistocene hominins from African, Asian and European sites in order to complete our analysis of the upper postcanine dentition. Intraspecific variation in these two molars is high, but some interspecific trends can be identified. Both molars exhibit a strong reduction of the distal cusps in recent hominin species, namely European Homo heidelbergensis, Homo neanderthalensis and Homo sapiens, but this reduction shows specific patterns and proportions in the three groups. Second molars tend to show four well developed cusps in earlier hominin species and their morphology is only marginally affected by allometric effects. Third molars can be incipiently reduced in earlier species and they evince a significant allometric component, identified both inter- and intraspecifically. European Middle Pleistocene fossils from Sima de los Huesos (SH) show a very strong reduction of these two molars, even more marked than the reduction observed in Neanderthals and in modern human populations. The highly derived shape of SH molars points to an early acquisition of typical Neanderthal dental traits by pre-Neanderthal populations and to a deviation of this population from mean morphologies of other European Middle Pleistocene groups.     

The Neanderthal face is not cold adapted

Many morphological features of the Pleistocene fossil hominin Homo neanderthalensis, including the reputed large size of its paranasal sinuses, have been interpreted as adaptations to extreme cold, as some Neanderthals lived in Europe during glacial periods. This interpretation of sinus evolution rests on two assumptions: that increased craniofacial pneumatization is an adaptation to lower ambient temperatures, and that Neanderthals have relatively large sinuses. Analysis of humans, other primates, and rodents, however, suggests that the first assumption is suspect; at least the maxillary sinus undergoes a significant reduction in volume in extreme cold, in both wild and laboratory conditions. The second assumption, that Neanderthal sinuses are large, extensive, or even ‘hyperpneumatized,’ has held sway since the first specimen was described and has been interpreted as the causal explanation for some of the distinctive aspects of Neanderthal facial form, but has never been evaluated with respect to scaling. To test the latter assumption, previously published measurements from two-dimensional (2D) X-rays and new three-dimensional (3D) data from computed tomography (CT) of Neanderthals and temperate-climate European Homo sapiens are regressed against cranial size to determine the relative size of their sinuses. The 2D data reveal a degree of craniofacial pneumatization in Neanderthals that is both commensurate with the size of the cranium and comparable in scale with that seen in temperate climate H. sapiens. The 3D analysis of CT data from a smaller sample supports this conclusion. These results suggest that the distinctive Neanderthal face cannot be interpreted as a direct result of increased pneumatization, nor is it likely to be an adaptation to resist cold stress; an alternative explanation is thus required.     

Homo erectus found still further west: Reconstruction of the Kocabaş cranium (Denizli,Turkey)

Few human fossils are known in Turkey and no Homo erectus has been discovered until now. In this respect, the newly discovered partial skull from Kocaba? is very important: (1) to assess the pattern of the first settlements throughout the Old World; and (2) to document the extension of the species H. erectus to the west of continental Asia. Using CT data and 3D imaging techniques, this specimen was reconstructed and a more detailed analysis was done, including the inner anatomical features. The preliminary results of this study highlight that the fossil hominid from Kocaba? is close to the H. erectus species regarding the following cranial patterns: presence of a clear post-orbital constriction, strong development of the frontal brow-ridge with a depressed supratoral area in the lateral part, as well as endocranial patterns such as the development and orientation of the middle meningeal artery and the presence of a frontal bec. The Kocaba? skull is morphologically very close to the fossils from Zhoukoudian L-C. The partial Kocaba? skull is the oldest H. erectus known in Turkey and the only one from this species to have settled so far west in Asia.     

安徽巢县人类化石地点的新材料

1983年对巢县银山人类化石地点的发掘结果再次表明,该地点的人类化石的地质时代大致相当于北京猿人地点1—4层的或稍晚。发现人类上颌骨化石一块,具有早期智人的特征。     

Variations and asymmetries in regional brain surface in the genus Homo

Paleoneurology is an important field of research within human evolution studies. Variations in size and shape of an endocast help to differentiate among fossil hominin species whereas endocranial asymmetries are related to behavior and cognitive function. Here we analyse variations of the surface of the frontal, parieto-temporal and occipital lobes among different species of Homo, including 39 fossil hominins, ten fossil anatomically modern Homo sapiens and 100 endocasts of extant modern humans. We also test for the possible asymmetries of these features in a large sample of modern humans and observe individual particularities in the fossil specimens.This study contributes important new information about the brain evolution in the genus Homo. Our results show that the general pattern of surface asymmetry for the different regional brain surfaces in fossil species of Homo does not seem to be different from the pattern described in a large sample of anatomically modern H. sapiens, i.e., the right hemisphere has a larger surface than the left, as do the right frontal, the right parieto-temporal and the left occipital lobes compared with the contra-lateral side. It also appears that Asian Homo erectus specimens are discriminated from all other samples of Homo, including African and Georgian specimens that are also sometimes included in that taxon. The Asian fossils show a significantly smaller relative size of the parietal and temporal lobes. Neandertals and anatomically modern H. sapiens, who share the largest endocranial volume of all hominins, show differences when considering the relative contribution of the frontal, parieto-temporal and occipital lobes. These results illustrate an original variation in the pattern of brain organization in hominins independent of variations in total size. The globularization of the brain and the enlargement of the parietal lobes could be considered derived features observed uniquely in anatomically modern H. sapiens.     

Out of the North Sea: the Zeeland Ridges Neandertal

In 2001, a portion of human frontal bone was discovered in sediments extracted from the bottom of the North Sea, 15 km off the coast of the Netherlands. The extraction zone is located in the so-called Zeeland Ridges area located at 51°40′ northern latitude and 3°20′ eastern longitude. The specimen was dredged up from sediments containing Late Pleistocene faunal remains and Middle Palaeolithic artefacts, including well-finished small handaxes and Levallois flakes. The details of the supraorbital morphology, as well as the quantitative assessment of the shape of the external surface of the squama using traditional and 3D geometric morphometrics, unambiguously assign the Zeeland Ridges frontal bone to Homo neanderthalensis. Carbon and nitrogen isotopic analysis indicate that the Zeeland Ridges hominin, like other Neandertals, was highly carnivorous and does not show evidence for the consumption of aquatic foods. A lesion on the outer table and diploic layer of the bone in the area of the supratoral sulcus can be interpreted as the result of an intradiploic epidermoid cyst, a type of neoplasm diagnosed for the first time in Neandertal remains. So far, the Zeeland Ridges Neandertal is the first Pleistocene fossil hominin found under seawater and the first recorded in the Netherlands.     

安徽和县猿人化石及有关问题的初步研究

安徽省和县猿人化石产地龙潭洞自1974至1980,先后经和县文化局、省水文地质工程地质队、省文物局和中国科学院古脊椎动物与古人类研究所等单位的调查和发掘,获得了一个猿人头盖骨,部分下颌骨和零星牙齿。这些材料大约代表3个个体,包括青年、壮年和老年。在同一层位发现的脊椎动物化石经初步鉴定,大约50种。另外,在遗址中还发现一些骨、角制品和烧过的骨、牙碎片。化石层的地质时代属更新世中期,昔日的生态环境为森林草原,气候趋于凉爽。     

陕西黄龙出土的人类头盖骨化石

黄龙人头盖骨是1975年在陕西省黄龙县徐家坟山的南坡挖到的。它出土于黄土层与红色土的交界处,从地层上看,可能属于更新世晚期。 该头盖骨保留了额骨和顶骨部分。从颅穹隆起程度、眶上区和颞区发育程度以及额鳞倾斜程度来判断,它属于晚期智人阶段。与晚期智人一般特征相比,它的矢状脊较突出,头骨骨壁较厚,额结节较不发育,前囟点位置较靠后。黄龙人头盖骨上现代特征和较原始特征的混合表明它代表了晚期智人中的一种过渡性的类型。     

L’architecture de l’épaule au sein du genre Homo : nouvelles interprétations

The morphology of human clavicles can be estimated by projecting them on two perpendicular planes in order to assess the shapes of their cranial and dorsal primary curvatures. In cranial view no differences in curvature appear within the genus Homo, which means the different species had similar arms elevation capacity, especially in protraction. On the contrary, in dorsal view two clavicles morphologies could be defined. The first one is characterized by two curvatures in dorsal view and is possessed by all Homo species, from Homo habilis to Neanderthal, including Homo ergaster, but not modern human, Upper Paleolithic and anatomically modern human remains, who possess clavicles of the second type, characterized by either one curvature, or two slightly pronounced ones in dorsal view. Clavicles displaying two pronounced curvatures in dorsal view are associated with scapula sitting high on the thorax in regard to modern human. However, shoulder with high scapula on the thorax displays two different kinds of architectures: (i) shoulder with short clavicles associated to scapulas sitting more laterally than those of modern human. This group includes earlier Homo like Homo habilis and Homo ergaster and (ii) shoulder with long clavicles associated to scapulas sitting more dorsally on the thorax, like those of modern human. This group includes Homoantecessor and Neanderthals. In other words, within the genus Homo, three shoulders would have existed. Evolution of the shoulder complex is far more complex than previously thought and the arrival of modern bipedalism was not associated to modern shoulder.