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The analysis of a large sample of anthracothere remains discovered by the Mission Paléoanthropologique Franco-Tchadienne at Toros-Menalla (TM), Northern Chad, has revealed the occurrence of a single species with a large intraspecific morphological variability. Taxonomic problems triggered the necessity of naming a new species for the TM anthracotheres,Libycosaurus bahrisp. nov. The description of the craniodental remains and the analysis of their variability indicate a marked sexual dimorphism, most probably induced by sex differential growth, helping to reassess the specific diversity ofLibycosaurusand its main evolutionary trends in a phylogenetic perspective. The ecological niche occupied by the new species was also further characterized in its semiaquatic, behavioural, and dietary dimensions. On these grounds we suggest that Late Miocene environmental changes, particularly habitat opening and hydrographic network fragmentation related to increased aridity, played a major role in canalizing semi-aquatic specializations and the extinction of the last African anthracotheres, as well as other bothriodontines. http://zoobank.org/urn:lsid:zoobank.org:pub:8C0AD4BD-952D-4A60-94B8-6F7A23332171 [ABSTRACT FROM AUTHOR]

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.

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., (Copyright © 2014 Elsevier Ltd. All rights reserved.)

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.

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.

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.

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.

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.

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.

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.

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.

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.

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.

All six known specimens of the early hominid Sahelanthropus tchadensis come from Toros-Menalla site 266 (TM 266), a single locality in the Djurab Desert, northern Chad, central Africa. Here we present a preliminary analysis of the palaeontological and palaeoecological context of these finds. The rich fauna from TM 266 includes a significant aquatic component such as fish, crocodiles and amphibious mammals, alongside animals associated with gallery forest and savannah, such as primates, rodents, elephants, equids and bovids. The fauna suggests a biochronological age between 6 and 7 million years. Taken together with the sedimentological evidence, the fauna suggests that S. tchadensis lived close to a lake, but not far from a sandy desert, perhaps the oldest record of desert conditions in the Neogene of northern central Africa.

The search for the earliest fossil evidence of the human lineage has been concentrated in East Africa. Here we report the discovery of six hominid specimens from Chad, central Africa, 2,500 km from the East African Rift Valley. The fossils include a nearly complete cranium and fragmentary lower jaws. The associated fauna suggest the fossils are between 6 and 7 million years old. The fossils display a unique mosaic of primitive and derived characters, and constitute a new genus and species of hominid. The distance from the Rift Valley, and the great antiquity of the fossils, suggest that the earliest members of the hominid clade were more widely distributed than has been thought, and that the divergence between the human and chimpanzee lineages was earlier than indicated by most molecular studies.

Abel is the first Australopithecine known west of the East African Rift Valley. The fossil finds include a perisymphyseal fragment of an adult mandible with well-preserved crowns of the right I2-P4 and the left C-P4. Abel's dentition displays many enamel defects, which are described in detail for each tooth. Pitting affected every tooth, while larger, shallow depressions were observed on the canines alone. From two to four pit bands occurred on the different teeth, each resulting from a separate hypoplastic episode. In modern humans, a large number of causes, both environmental and genetic, have been suggested for such enamel defects. It seems probable that Abel's pathology was systemic. However, the occurrence of a number of bands and the variable intensity of the attacks make it difficult to say whether there was a single, repetitive cause or different etiologies. Possible causes discussed here include recurrent fevers and seasonal nutritional deficiencies.

The first sites with Pliocene and Pleistocene mammals west of the Rift Valley in Central Africa in northern Chad were reported in 1959 (ref. 1), and documented the presence of mixed savannah and woodland habitats. Further sites and a probable Homo erectus cranio-facial fragment were subsequently discovered. In 1993 a survey of Pliocene and Pleistocene formations in the Borkou-Ennedi-Tibesti Province of Chad (B.E.T.) led to the discovery of 17 new sites in the region of Bahr el Ghazal (classical Arabic for River of the Gazelles) near Koro Toro. One site, KT 12 (15 degrees 58'10"N, 18 degrees 52'46"E) yielded an australopithecine mandible associated with a fauna biochronologically estimated to be 3.0-3.5 Myr old. Australopithecine species described since 1925 are known from southern Africa and from sites spread along the eastern Rift Valley from Tanzania to Ethiopia (Fig. 1). This new find from Chad, which is most similar in morphology to Australopithecus afarensis, documents the presence of an early hominid a considerable distance, 2,500 km, west of the Rift Valley.