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.

Relationship between foramen magnum position and locomotion in extant and extinct hominoids.

From the Miocene Sahelanthropus tchadensis to Pleistocene Homo sapiens, hominins are characterized by a derived anterior position of the foramen magnum relative to basicranial structures. It has been previously suggested that the anterior position of the foramen magnum in hominins is related to bipedal locomotor behavior. Yet, the functional relationship between foramen magnum position and bipedal locomotion remains unclear. Recent studies, using ratios based on cranial linear measurements, have found a link between the anterior position of the foramen magnum and bipedalism in several mammalian clades: marsupials, rodents, and primates. In the present study, we compute these ratios in a sample including a more comprehensive dataset of extant hominoids and fossil hominins. First, we verify if the values of ratios can distinguish extant humans from apes. Then, we test whether extinct hominins can be distinguished from non-bipedal extant hominoids. Finally, we assess if the studied ratios are effective predictors of bipedal behavior by testing if they mainly relate to variation in foramen magnum position rather than changes in other cranial structures. Our results confirm that the ratios discriminate between extant bipeds and non-bipeds. However, the only ratio clearly discriminating between fossil hominins and other extant apes is that which only includes basicranial structures. We show that a large proportion of the interspecific variation in the other ratios relates to changes in facial, rather than basicranial, structures. In this context, we advocate the use of measurements based only on basicranial structures when assessing the relationship between foramen magnum position and bipedalism in future studies.

New data about semicircular canal morphology and locomotion in modern hominoids.

The labyrinth has two functional parts: the cochlea for audition and the vestibular system for equilibrioception. In the latter, the semicircular ducts and the otolithic organs are sensitive to rotational and linear accelerations of the head, respectively. The labyrinthine morphology influences perception accuracy, hence the adaptation to a specific locomotor pattern. The aim of this study is to determine the relationship between locomotion and semicircular canal morphology using geometric morphometrics, and to explain these links with existing functional models. The influence of factors other than functional constraints on labyrinthine morphology is discussed. The left bony labyrinth of 65 specimens was extracted virtually. Five extant hominoid species with various locomotion modes were sampled. A set of 13 landmarks was placed on the semicircular canals. After a Procrustes fit, their coordinates were analyzed using a principal component analysis. It was found that labyrinthine morphology is significantly distinct between species. More specifically, the differences involve a posterolateral projection of the lateral semicircular canal and the rotation of this canal relative to the vertical canals. This rotation occurs in the sagittal plane, which is consistent with previous studies based on traditional morphometrics. Among extant hominoids, the shape of the canals potentially discriminates species based on posture. This result could be used to reconstruct the locomotor pattern of fossil hominoids.

Phytoliths indicate significant arboreal cover at Sahelanthropus type locality TM266 in northern Chad and a decrease in later sites.

We analyzed phytolith and diatom remains preserved at 45 Miocene and Pliocene localities dated between 8 and 1 Ma in northern Chad (16-17°N). Some of these localities yielded cranial remains, lower jaws, and teeth of the hominin species Australopithecus bahrelghazali (∼3.6 Ma) and Sahelanthropus tchadensis (∼7 Ma). Of the 111 sediment samples analyzed, 41 yielded phytoliths, 20 yielded diatoms, and seven yielded both phytoliths and diatoms. Freshwater planktonic and tychoplanktonic diatom species, indicative of lacustrine conditions, are dominant (>91%) in the samples. The phytolith assemblages indicate an opening of the vegetation and a general trend toward an expansion of grass-dominated environments during the time spanning the two hominin occurrences in Chad. The phytoliths suggest the presence of a mosaic environment, including closed forest patches, palm groves, and mixed/grassland formations, between 7.5 and 7 Ma, the replacement by palm grove-like vegetation at approximately 6.5-5 Ma, and the presence of exclusive grass-dominated formations after 4.5 Ma. The type-locality of S. tchadensis (TM266) was likely similar to modern palm grove formations with an arboreal cover percentage ≥40%. The type locality of A. bahrelghazali (KT12) was a grass-dominated ecosystem (likely savanna) with an unrated percentage of arboreal cover. Furthermore, the grass phytolith data support the existence of a (recurrent) Sahelian-like dry climate in northern Chad since at least 8 Ma. Therefore the local closed vegetation formations in the Djurab region at 7.5-7 Ma were sustained by aquatic systems (such as lakes or related rivers, marshes) rather than by extensive annual precipitation.

A new species of Nyanzachoerus (Cetartiodactyla: Suidae) from the late Miocene Toros-Ménalla, Chad, central Africa.

During the latest Miocene and the early Pliocene, tetraconodontine suids were the most predominant large omnivorous mammals in Africa. Yet, new species were often identified on the grounds of limited evidence, a situation impacting their value for biochronological correlations as well as for environmental and biogeographical reconstructions. The description of the most abundant known collection of craniodental remains attributed to the tetraconodontine Nyanzachoerus helps to improve this situation. These specimens were collected in the upper Miocene deposits at Toros-Ménalla, northern Chad, central Africa, by the Mission Paléoanthropologique Franco-Tchadienne. We compared them with Nyanzachoerus from eastern and southern Africa, using extant species as a reference for patterns of morphological variation. Thanks to a large sample of observations, our work focused as much on craniomandibular morphology as on dental morphology and metrics (improved by an index scoring for the complexity of distal third molars and a detailed investigation of premolar-molar ratios). We recognized two taxa at Toros-Ménalla: Nyanzachoerus khinzir nov. sp. and Ny. cf. australis. We also revised the taxonomic status for other species, including: the restriction of Ny. syrticus to its holotype specimen from Sahabi (Libya), the resurrection of the nomen Ny. tulotos, and the synonymy of Ny. kuseralensis with Ny. waylandi. At Toros-Ménalla, Ny. khinzir was the only suid coexisting with the anthracotheriid Libycosaurus and the hominid Sahelanthropus, whereas Ny. cf. australis was associated with a different, probably younger faunal context. Nyanzachoerus. khinzir, which probably had a diversified diet, supports a latest Miocene biogeographical distinction between central Africa and eastern Africa.

Toros-Menalla (Chad, 7 Ma), the earliest hominin-bearing area: How many mammal paleocommunities?

The fossiliferous area of Toros-Menalla (TM) (Djurab Desert, northern Chad) has yielded one of the richest African mammal faunas of the late Miocene. It is also the place where the earliest known hominin, Sahelanthropus tchadensis, was found. Although more than 300 localities are recorded in that area, previous paleoecological studies focused only on the largest and richest one. The integration of the material from other TM localities, and thus of a significant number of mammal taxa, is crucial to improve the corresponding paleoenvironmental reconstructions. Before such inferences can be drawn, it is necessary to test for the ecological integrity of these mammal assemblages: how many paleocommunities do they represent? The faunal structures of several assemblages selected for their apparent resilience to sampling biases are compared here. The criteria used in the inter-assemblage comparison are ecological diversity, taxonomic structure (taxonomic rank of abundance) and taxonomic composition. Based on multivariate analyses, two groups of TM assemblages can be distinguished. One of them contains the hominin-bearing assemblages. It is taxonomically richer and shows a wider ecological spectrum than its counterpart. The degree of taphonomic alteration undergone by the TM assemblages, as well as the distribution of amphibious mammals among them, suggest different depositional settings for these two groups of assemblages, the richest of which was probably associated with lower hydraulic energy. Overall, it seems that the TM assemblages recorded the same mammal paleocommunity preserved in two contrasted depositional settings. Moreover, the spatial overlap of these assemblages provides further evidence for the mosaic character of the landscape associated with S. tchadensis.

Subocclusal dental morphology of sahelanthropus tchadensis and the evolution of teeth in hominins.

The evolution of the teeth in hominins is characterized by, among other characters, major changes in root morphology. However, little is known of the evolution from a plesiomorphic, ape-like root morphology to the crown hominin morphology. Here we present a study of the root morphology of the Miocene Chadian hominin Sahelanthropus tchadensis and its comparison to other hominins. The morphology of the whole lower dentition (I1 -M3 ) was investigated and described. The comparison with the species Ardipithecus kaddaba and Ardipithecus ramidus indicates a global homogeneity of root morphology in early hominins. This morphology, characterized notably by a reduction of the size and number of the roots of premolars, is a composite between an ape-like morphology and the later hominin morphology. Trends for root evolution in hominins are proposed, including the transition from a basal hominoid to extant Homo sapiens. This study also illustrates the low association between the evolution of tooth root morphology and the evolution of crowns in hominins.

New Apterodontinae (Hyaenodontida) from the Eocene locality of Dur At-Talah (Libya): systematic, paleoecological and phylogenetical implications.

The African Hyaenodontida, mainly known from the Late Eocene and Early Oligocene Fayum depression in Egypt, show a very poor diversity in oldest Paleogene localities. Here we report new hyaenodontidans found in the late Middle Eocene deposits of Dur At-Talah (Central Libya), known to have recorded the earliest radiation of African anthropoids. The new hyaenodontidan remains are represented by dental and postcranial specimens comprising the historical material discovered by R.J.G. Savage in the last century and that of the recent Franco-Libyan campaigns. This material includes two apterodontines, in particular a subcomplete skeleton of Apterodon langebadreae nov. sp., bringing new postcranial elements to the fossil record of the genus Apterodon. Anatomical analysis of the postcranial remains of Dur At-Talah suggests a semi-aquatic lifestyle for Apterodon, a completely unusual locomotion pattern among hyaenodontidans. We also perform the first cladistic analysis of hyaenodontidans including apterodontines: Apterodon and Quasiapterodon appear close relatives to "hyainailourines", in particular to the African Oligo-Miocene Metasinopa species. Apterodon langebadreae nov. sp. could be the most primitive species of the genus, confirming an African origin of the Apterodontinae and a further dispersion event to Europe before the early Oligocene. These data enhance our knowledge of early hyaenodontidan diversification into Africa and underline how crucial is the understanding of their evolutionary history for the improvement of Paleogene paleobiogeographic scenarii.

Isotopic evidence for an early shift to C4 resources by Pliocene hominins in Chad.

Foods derived from C(4) plants were important in the dietary ecology of early Pleistocene hominins in southern and eastern Africa, but the origins and geographic variability of this relationship remain unknown. Carbon isotope data show that Australopithecus bahrelghazali individuals from Koro Toro in Chad are significantly enriched in (13)C, indicating a dependence on C(4) resources. As these sites are over 3 million years in age, the results extend the pattern of C(4) dependence seen in Paranthropus boisei in East Africa by more than 1.5 million years. The Koro Toro hominin fossils were found in argillaceous sandstone levels along with abundant grazing and aquatic faunal elements that, in combination, indicate the presence of open to wooded grasslands and stream channels associated with a greatly enlarged Lake Chad. In such an environment, the most abundant C(4) plant resources available to A. bahrelghazali were grasses and sedges, neither of which is usually considered as standard great ape fare. The results suggest an early and fundamental shift in hominin dietary ecology that facilitated the exploitation of new habitats.

Assessing endocranial variations in great apes and humans using 3D data from virtual endocasts.

Modern humans are characterized by their large, complex, and specialized brain. Human brain evolution can be addressed through direct evidence provided by fossil hominid endocasts (i.e. paleoneurology), or through indirect evidence of extant species comparative neurology. Here we use the second approach, providing an extant comparative framework for hominid paleoneurological studies. We explore endocranial size and shape differences among great apes and humans, as well as between sexes. We virtually extracted 72 endocasts, sampling all extant great ape species and modern humans, and digitized 37 landmarks on each for 3D generalized Procrustes analysis. All species can be differentiated by their endocranial shape. Among great apes, endocranial shapes vary from short (orangutans) to long (gorillas), perhaps in relation to different facial orientations. Endocranial shape differences among African apes are partly allometric. Major endocranial traits distinguishing humans from great apes are endocranial globularity, reflecting neurological reorganization, and features linked to structural responses to posture and bipedal locomotion. Human endocasts are also characterized by posterior location of foramina rotunda relative to optic canals, which could be correlated to lesser subnasal prognathism compared to living great apes. Species with larger brains (gorillas and humans) display greater sexual dimorphism in endocranial size, while sexual dimorphism in endocranial shape is restricted to gorillas, differences between males and females being at least partly due to allometry. Our study of endocranial variations in extant great apes and humans provides a new comparative dataset for studies of fossil hominid endocasts.

Late middle Eocene epoch of Libya yields earliest known radiation of African anthropoids.

Reconstructing the early evolutionary history of anthropoid primates is hindered by a lack of consensus on both the timing and biogeography of anthropoid origins. Some prefer an ancient (Cretaceous) origin for anthropoids in Africa or some other Gondwanan landmass, whereas others advocate a more recent (early Cenozoic) origin for anthropoids in Asia, with subsequent dispersal of one or more early anthropoid taxa to Africa. The oldest undoubted African anthropoid primates described so far are three species of the parapithecid Biretia from the late middle Eocene Bir El Ater locality of Algeria and the late Eocene BQ-2 site in the Fayum region of northern Egypt. Here we report the discovery of the oldest known diverse assemblage of African anthropoids from the late middle Eocene Dur At-Talah escarpment in central Libya. The primate assemblage from Dur At-Talah includes diminutive species pertaining to three higher-level anthropoid clades (Afrotarsiidae, Parapithecidae and Oligopithecidae) as well as a small species of the early strepsirhine primate Karanisia. The high taxonomic diversity of anthropoids at Dur At-Talah indicates either a much longer interval of anthropoid evolution in Africa than is currently documented in the fossil record or the nearly synchronous colonization of Africa by multiple anthropoid clades at some time during the middle Eocene epoch.

Two new Mio-Pliocene Chadian hominids enlighten Charles Darwin's 1871 prediction.

The idea of an evolutionary sequence for humans is quite recent. Over the last 150 years, we have discovered unexpected ancestors, numerous close relatives and our deep evolutionary roots in Africa. In the last decade, three Late Miocene hominids have been described, two about 6 Ma (Ardipithecus and Orrorin) in East Africa and the third dated to about 7 Ma (Sahelanthropus) in Central Africa. The specimens are too few to propose definite relationship to other species, but clearly these belong to a new evolutive grade distinct from Australopithecus and Homo. Moreover, all of them were probably habitual bipeds and lived in woodlands, thus falsifying the savannah hypothesis of human origins. In light of all this recent knowledge, Charles Darwin predicted correctly in 1871 that Africa is the birthplace of humans, chimpanzees and our close relatives.

Gaudeamus lavocati sp. nov. (Rodentia, Hystricognathi) from the early Oligocene of Zallah, Libya: first African caviomorph?

A new African species of hystricognathous rodent, Gaudeamus lavocati sp. nov., is described herein from the early Oligocene deposits of Zallah locality (Sirt basin, Central Libya). The dental morphology of this species is very close to that of some earliest South American caviomorphs. It allows a reinterpretation of molar crest homologies among earliest caviomorphs, pentalophodonty being confirmed as the plesiomorphic molar condition in Caviomorpha. This morphological resemblance argues for close affinities between Gaudeamus and earliest South American hystricognaths. Cladistic analysis supports Gaudeamus lavocati sp. nov. as the first known African representative of Caviomorpha, implying that its ancestors were part of the African phiomyid group that crossed the South Atlantic by a direct immigration route. Alternatively, the series of derived dental features of Gaudeamus could also be interpreted as evolutionary synchronous convergences of an African hystricognath lineage towards the specialized pattern of some caviomorphs. However, the high level of similarities concerning teeth morphology and enamel microstructure and the similar age of fossiliferous strata on both continents make this interpretation less probable. The phylogenetic position of this taxon is of considerable importance because it represents an enigmatic component of the phiomorph-caviomorph radiation in Africa and appears as a new clue toward the understanding of caviomorph origins.

Macrovertebrate paleontology and the Pliocene habitat of Ardipithecus ramidus.

A diverse assemblage of large mammals is spatially and stratigraphically associated with Ardipithecus ramidus at Aramis. The most common species are tragelaphine antelope and colobine monkeys. Analyses of their postcranial remains situate them in a closed habitat. Assessment of dental mesowear, microwear, and stable isotopes from these and a wider range of abundant associated larger mammals indicates that the local habitat at Aramis was predominantly woodland. The Ar. ramidus enamel isotope values indicate a minimal C4 vegetation component in its diet (plants using the C4 photosynthetic pathway), which is consistent with predominantly forest/woodland feeding. Although the Early Pliocene Afar included a range of environments, and the local environment at Aramis and its vicinity ranged from forests to wooded grasslands, the integration of available physical and biological evidence establishes Ar. ramidus as a denizen of the closed habitats along this continuum.

The mammal assemblage of the hominid site TM266 (Late Miocene, Chad Basin): ecological structure and paleoenvironmental implications.

Characterizing the paleoenvironmental context of the first hominids is a key issue for understanding their behavioral and morphological evolution. The present study aims at reconstructing the paleoenvironment of the TM266 vertebrate assemblage (Toros-Menalla, Northern Chad) that yielded the earliest known hominid Sahelanthropus tchadensis (7 Ma). For the first time, a quantitative analysis is carried out on the fossil mammal assemblage associated with that hominid. Two complementary approaches were applied: (1) the analysis of the relative abundances of taxa and their habitat preferences; and (2) the study of the distribution of taxa within three meaningful ecovariables: locomotion, feeding preferences, and body mass. The resulting taxonomic and paleoecological structures are used to reconstruct the diversity and the relative extent of the habitats in that part of northern Chad seven million years ago. The paleoenvironment was composed of open areas with dry and humid grasslands, prevailing over wooded habitats. Water was also widely available as freshwater bodies and certainly swamps. It appears that the high habitat diversity of the landscape is a common feature among paleoenvironments associated with early hominids.

Analysis of type I endoleaks in a stented abdominal aortic aneurysm.

To improve the performance of endovascular grafts used for the treatment of abdominal aortic aneurysms, we develop a methodology to analyze the phenomena of type I endoleaks in a non-invasive-stented abdominal aorta. As one aspect of this study, an evaluation of the parietal stresses generated by the blood flow is provided. As blood is known to be a shear-thinning, non-Newtonian fluid, we have chosen to use the Phan-Thien and Tanner model, which can be derived from the rheology of polymer solutions. As a second aspect, we develop an axisymmetric finite-element model of the complete system. An explicit finite-element in-house code, is used to simulate the behavior of the system, which is subjected to hydrostatic pressure and to the stresses generated by the blood flow. As the response of the solid is strongly affected by the response of the fluid, and vice versa, the modeling of a coupled fluid-structure interaction is achieved in this work. This study provides an evaluation of the stresses generated by the blood flow on the aorta's wall. The finite-element model allows to identify biomechanical factors that can influence the propensity of an aneurysm treated with an endograft, to exhibit endoleaks. First observations are made concerning the influence of oversizing of the endograft and the influence of friction coefficients between the aorta, endograft and plaque.

Cosmogenic nuclide dating of Sahelanthropus tchadensis and Australopithecus bahrelghazali: Mio-Pliocene hominids from Chad.

Ages were determined at two hominid localities from the Chad Basin in the Djurab Desert (Northern Chad). In the Koro Toro fossiliferous area, KT 12 locality (16 degrees 00'N, 18 degrees 53'E) was the site of discovery of Australopithecus bahrelghazali (Abel) and in the Toros-Menalla fossiliferous area, TM 266 locality (16 degrees 15'N, 17 degrees 29'E) was the site of discovery of Sahelanthropus tchadensis (Toumaï). At both localities, the evolutive degree of the associated fossil mammal assemblages allowed a biochronological estimation of the hominid remains: early Pliocene (3-3.5 Ma) at KT 12 and late Miocene ( approximately 7 Ma) at TM 266. Atmospheric (10)Be, a cosmogenic nuclide, was used to quasicontinuously date these sedimentary units. The authigenic (10)Be/(9)Be dating of a pelite relic within the sedimentary level containing Abel yields an age of 3.58 +/- 0.27 Ma that points to the contemporaneity of Australopithecus bahrelghazali (Abel) with Australopithecus afarensis (Lucy). The 28 (10)Be/(9)Be ages obtained within the anthracotheriid unit containing Toumaï bracket, by absolute dating, the age of Sahelanthropus tchadensis to lie between 6.8 and 7.2 Ma. This chronological constraint is an important cornerstone both for establishing the earliest stages of hominid evolution and for new calibrations of the molecular clock.

Symphyseal shape variation in extant and fossil hominoids, and the symphysis of Australopithecus bahrelghazali.

The holotype of the species Australopithecus bahrelghazali is a mandibular fragment preserving left C-P(4) and right I(2)-P(4). One of the key features of the A. bahrelghazali mandible is its sagittally and transversally flat anterior region associated with a vertical, bulbous symphysis that is assumed to differ morphologically from the classic, more apelike eastern australopith morphology with its sloping symphysis, developed transverse tori, and distinct genioglossal fossa. Yet, close similarity has been suggested between A. bahrelghazali and A. afarensis. To date, no metrical comparison of the symphyseal morphology of east and west African australopiths has been performed. For the selected characters, this study attempts to test the following null hypothesis: A. bahrelghazali does not present significant differences from A. afarensis (i.e., A. bahrelghazali vs. A. afarensis variation does not depart from expected intraspecific variation for hominoid species). Analysis of the mandibular symphysis is difficult to undertake using conventional linear variables since few landmarks are available to make a precise quantitative assessment of its morphology. In addition, while a few studies have used outline-based techniques of analysis to address symphyseal shape variation, none has integrated data on early hominins. We present here a detailed comparative study of variation in the symphyseal outline of A. bahrelghazali and A. afarensis based on a quantitative characterization by elliptic Fourier coefficients. Original data on symphyseal variation in Pliocene hominins are provided and discussed within a comparative framework of extant and fossil hominoid representatives. We evaluate the relevance of our quantitative data describing the shape of the symphysis for discriminating hominoid taxa, and test for differences in symphyseal shape between A. bahrelghazali and A. afarensis. Elliptic Fourier coefficients appear to be well-suited descriptors for depicting symphyseal variation within hominoids. Our results confirm that symphyseal shape is a good indicator of taxonomic affinity within hominoids, enabling a clear distinction between great apes, early hominins, and modern humans. Furthermore, our results substantiate the unusual pattern of the A. bahrelghazali symphyses compared to A. afarensis and support the specific status of the west African australopith.

The oldest African fox (Vulpes riffautae n. sp., Canidae, Carnivora) recovered in late Miocene deposits of the Djurab desert, Chad.

We report on the oldest fox (Canidae) ever found in Africa. It is dated to 7 Ma based on the degree of evolution of the whole fauna. It belongs to a new species. Its overall size and some morphological characteristics distinguish the Chadian specimen from all the other foxes. The presence of Vulpes and of the genus Eucyon in slightly younger African locality, as well as in southwestern Europe in the late Miocene, may indicate that canids migrated in Europe from Africa through a trans-Mediterranean route.

Tectonic uplift and Eastern Africa aridification.

The history of Eastern African hominids has been linked to a progressive increase of open grassland during the past 8 million years. This trend was explained by global climatic processes, which do not account for the massive uplift of eastern African topography that occurred during this period. Atmosphere and biosphere simulations quantify the role played by these tectonic events. The reduced topographic barrier before 8 million years ago permitted a zonal circulation with associated moisture transport and strong precipitation. Our results suggest that the uplift itself led to a drastic reorganization of atmospheric circulation, engendering the strong aridification and paleoenvironmental changes suggested by the data.