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.

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.

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.

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.

The earliest modern mongoose (Carnivora, Herpestidae) from Africa (late Miocene of Chad).

We report on the earliest modern mongooses of Africa, from the late Miocene (ca. 7 Ma) of the hominid locality TM 266, Toros-Menalla, Chad. The material is based on fragmentary dentitions of three individuals. The main diagnostic feature of the Chadian species is the great development of the shear in the carnassials, which distinguishes the Chadian specimens from all extant herpestids except Herpestes and Galerella. In comparison with most extinct and extant Herpestes, the species from Toros-Menalla differs by a markedly smaller size and, depending on the species, relatively more elongated carnassials, more transversely elongated M1 and more reduced p4. On the basis of a great morphological similarity and the absence of significant differences, we assign our material to Galerella sanguinea; the Chadian finding therefore represents the earliest appearance of an extant species of Herpestidae. This record ties the first appearance of the genus to a minimum age of ca. 7 Ma, which is consistent with the estimated divergence date of 11.4 Ma known from the literature for the species of Galerella.

Geology and palaeontology of the Upper Miocene Toros-Menalla hominid locality, Chad.

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.

A new hominid from the Upper Miocene of Chad, 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.