Inner ear sensory system changes as extinct crocodylomorphs transitioned from land to water.

Major evolutionary transitions, in which animals develop new body plans and adapt to dramatically new habitats and lifestyles, have punctuated the history of life. The origin of cetaceans from land-living mammals is among the most famous of these events. Much earlier, during the Mesozoic Era, many reptile groups also moved from land to water, but these transitions are more poorly understood. We use computed tomography to study changes in the inner ear vestibular system, involved in sensing balance and equilibrium, as one of these groups, extinct crocodile relatives called thalattosuchians, transitioned from terrestrial ancestors into pelagic (open ocean) swimmers. We find that the morphology of the vestibular system corresponds to habitat, with pelagic thalattosuchians exhibiting a more compact labyrinth with wider semicircular canal diameters and an enlarged vestibule, reminiscent of modified and miniaturized labyrinths of other marine reptiles and cetaceans. Pelagic thalattosuchians with modified inner ears were the culmination of an evolutionary trend with a long semiaquatic phase, and their pelagic vestibular systems appeared after the first changes to the postcranial skeleton that enhanced their ability to swim. This is strikingly different from cetaceans, which miniaturized their labyrinths soon after entering the water, without a prolonged semiaquatic stage. Thus, thalattosuchians and cetaceans became secondarily aquatic in different ways and at different paces, showing that there are different routes for the same type of transition.

A window into the early evolutionary history of Cercopithecidae: Late Miocene evidence from Chad, Central Africa.

Central Africa is known as a major center of diversification for extant Old World Monkeys (OWM) and yet has a poorly documented fossil record of monkeys. Here we report a new colobine monkey (Cercopithecoides bruneti sp. nov.) from the Central African hominin-bearing fossiliferous area of Toros-Menalla, Chad at ca. 7 Ma. In addition to filling a gap in the spatial and temporal record of early OWM evolutionary history, we assess the ecomorphological diversity of early OWM by providing evidence on the onset of a folivorous diet and a partial reacquisition of terrestrial locomotor habits among Miocene colobines. We also support the phylogenetic affinities of the genus Cercopithecoides among the stem group of the extant African colobine monkeys.

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.

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.

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.

New material of the earliest hominid from the Upper Miocene of Chad.

Discoveries in Chad by the Mission Paleoanthropologique Franco-Tchadienne have substantially changed our understanding of early human evolution in Africa. In particular, the TM 266 locality in the Toros-Menalla fossiliferous area yielded a nearly complete cranium (TM 266-01-60-1), a mandible, and several isolated teeth assigned to Sahelanthropus tchadensis and biochronologically dated to the late Miocene epoch (about 7 million years ago). Despite the relative completeness of the TM 266 cranium, there has been some controversy about its morphology and its status in the hominid clade. Here we describe new dental and mandibular specimens from three Toros-Menalla (Chad) fossiliferous localities (TM 247, TM 266 and TM 292) of the same age. This new material, including a lower canine consistent with a non-honing C/P3 complex, post-canine teeth with primitive root morphology and intermediate radial enamel thickness, is attributed to S. tchadensis. It expands the hypodigm of the species and provides additional anatomical characters that confirm the morphological differences between S. tchadensis and African apes. S. tchadensis presents several key derived features consistent with its position in the hominid clade close to the last common ancestor of chimpanzees and humans.

Virtual cranial reconstruction of Sahelanthropus tchadensis.

Previous research in Chad at the Toros-Menalla 266 fossiliferous locality (about 7 million years old) uncovered a nearly complete cranium (TM 266-01-60-1), three mandibular fragments and several isolated teeth attributed to Sahelanthropus tchadensis. Of this material, the cranium is especially important for testing hypotheses about the systematics and behavioural characteristics of this species, but is partly distorted from fracturing, displacement and plastic deformation. Here we present a detailed virtual reconstruction of the TM 266 cranium that corrects these distortions. The reconstruction confirms that S. tchadensis is a hominid and is not more closely related to the African great apes. Analysis of the basicranium further indicates that S. tchadensis might have been an upright biped, suggesting that bipedalism was present in the earliest known hominids, and probably arose soon after the divergence of the chimpanzee and human lineages.

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.

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.

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.

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.

Facial orientation and facial shape in extant great apes: a geometric morphometric analysis of covariation.

The organization of the bony face is complex, its morphology being influenced in part by the rest of the cranium. Characterizing the facial morphological variation and craniofacial covariation patterns in extant hominids is fundamental to the understanding of their evolutionary history. Numerous studies on hominid facial shape have proposed hypotheses concerning the relationship between the anterior facial shape, facial block orientation and basicranial flexion. In this study we test these hypotheses in a sample of adult specimens belonging to three extant hominid genera (Homo, Pan and Gorilla). Intraspecific variation and covariation patterns are analyzed using geometric morphometric methods and multivariate statistics, such as partial least squared on three-dimensional landmarks coordinates. Our results indicate significant intraspecific covariation between facial shape, facial block orientation and basicranial flexion. Hominids share similar characteristics in the relationship between anterior facial shape and facial block orientation. Modern humans exhibit a specific pattern in the covariation between anterior facial shape and basicranial flexion. This peculiar feature underscores the role of modern humans' highly-flexed basicranium in the overall integration of the cranium. Furthermore, our results are consistent with the hypothesis of a relationship between the reduction of the value of the cranial base angle and a downward rotation of the facial block in modern humans, and to a lesser extent in chimpanzees.

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.

The first fossil fungus gardens of Isoptera: oldest evidence of symbiotic termite fungiculture (Miocene, Chad basin).

Higher termites of the subfamily Macrotermitinae (fungus-growing termites) are known to build fungus gardens where a symbiotic fungus (Termitomyces sp.) is cultivated. The fungus grows on a substrate called fungus comb, a structure built with the termites' own faeces. Here we present the first fossil fungus combs ever found in the world. They were extracted from 7-million-year-old continental sandstone (Chad basin). Fossilized fungus combs have an ovoid morphology with a more or less flattened concave base and a characteristic general alveolar aspect. Under lens, they display a typical millimetre-scale pelletal structure. The latter, as well as the general shape and alveolar aspect, are similar to the morphology of fungus combs from extant fungus-growing termites.

The age of the Sahara desert.

In the Sahara region, the age of onset of the desert condition has been uncertain until now. Here we report on the discovery of 7,000,000-year-old eolian dune deposits from the northern Chad Basin. This geological archive is the oldest known evidence for desert occurrence in the Sahara.

Anthracothere dental anatomy reveals a late Miocene Chado-Libyan bioprovince.

Recent discovery of an abundant and diverse late Miocene fauna at Toros-Ménalla (Chad, central Africa) by the Mission Paléoanthropologique Franco-Tchadienne provides a unique opportunity to examine African faunal and hominid evolution relative to the early phases of the Saharan arid belt. This study presents evidence from an African Miocene anthracotheriid Libycosaurus, particularly well documented at Toros-Ménalla. Its remains reveal a large semiaquatic mammal that evolved an autapomorphic upper fifth premolar (extremely rare in Cenozoic mammals). The extra tooth appeared approximately 12 million years ago, probably in a small northern African population isolated by climate-driven fragmentation and alteration of the environments inhabited by these anthracotheriids [Flower, B. P. & Kennett, J. P. (1994) Palaeogeogr. Palaeoclimatol. Palaeoecol. 108, 537-555 and Zachos, J., Pagani, M., Sloan, L., Thomas, E. & Billups, K. (2001) Science 292, 686-693]. The semiaquatic niche of Libycosaurus, combined with the distribution and relationships of its late Miocene species, indicates that by the end of the Miocene, wet environments connected the Lake Chad Basin to the Libyan Sirt Basin, across what is now the Sahara desert.

Pliocene large-mammal assemblages from Northern Chad: sampling and ecological structure.

Numerous Pliocene large-mammal assemblages have been discovered in Chad over the last decade. They offer a unique opportunity to understand the settings in which important chapters of Hominid evolution took place in Central Africa. However, it is crucial to first investigate both sampling and taxonomic homogeneity for these Chadian assemblages because they occur over large sectors in a sandy desert that offers virtually no stratigraphic section. Using cluster analysis and ordination techniques, we show that the three Pliocene sectors from Chad are homogeneous and adequate sampling units. Previous stable isotope analyses on these assemblages have indicated that the environment became richer in C(4) plants between approximately 5.3 and 3.5-3 Ma. To test whether this environmental change has affected the structure of palaeo-communities, we assigned body mass, trophic and locomotor eco-variables to mammal species from the three sectors. Statistical analysis shows that the overall ecological structure of the assemblages is not linked with the opening of the plant cover, and eco-variables show no temporal trend from the oldest sector to the youngest. For example, there is no significant change in the relative diversity of grazing and browsing taxa, although mixed feeders are less diversified in the youngest sector than in the preceding one. This pattern apparently does not result from potential biases such as methodological artefacts or taphonomic imprint. Instead, it seems that local heterogeneous environmental factors have played a major role in shaping the ecological spectrum of Chadian mammal palaeo-communities during the Pliocene.