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.

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.

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.

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.

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.

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.

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 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.

Morphological affinities of the Sahelanthropus tchadensis (Late Miocene hominid from Chad) cranium.

The recent reconstruction of the Sahelanthropus tchadensis cranium (TM 266-01-60-1) provides an opportunity to examine in detail differences in cranial shape between this earliest-known hominid, African apes, and other hominid taxa. Here we compare the reconstruction of TM 266-01-60-1 with crania of African apes, humans, and several Pliocene hominids. The results not only confirm that TM 266-01-60-1 is a hominid but also reveal a unique mosaic of characters. The TM 266-01-60-1 reconstruction shares many primitive features with chimpanzees but overall is most similar to Australopithecus, particularly in the basicranium. However, TM 266-01-60-1 is distinctive in having the combination of a short subnasal region associated with a vertical upper face that projects substantially in front of the neurocranium. Further research is needed to determine the evolutionary relationships between Sahelanthropus and the known Miocene and Pliocene hominids.

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.

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.

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.

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.