Pauline Coster

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.

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.

A complete magnetic-polarity stratigraphy of the Miocene continental deposits of Mae Moh Basin, northern Thailand, and a reassessment of the age of hominoid-bearing localities in northern Thailand

Northern Thailand comprises more than 40 Tertiary intermontane basins. These basins, tectonically interpreted as grabens or half grabens, have yielded hominoid fossils and rich Neogene mammal faunas. Relative ages provided by biochronological studies are controversial and neither absolute ages nor correlations with marine deposits are available. A precise temporal framework provided by magnetic-polarity stratigraphy is therefore crucial to understand the evolution of these Southeast Asian mammalian faunas, to estimate the timing and duration of basin formation, and finally to allow correlations with other Neogene hominoid localities of the Old World. A complete magnetostratigraphic study, which significantly extends previous paleomagnetic investigations, has been conducted in the Miocene sequence of Mae Moh Basin, Lampang Province, northern Thailand. Based on the biostratigraphic constraints, the 15 polarity zones recognized from the composite section have been correlated with Chrons C5ACr–C5r.3r, between 14.1 and 12.0 Ma. The present study provides a high-resolution magnetostratigraphic reference for the continental middle Miocene of all Southeast Asia. By correlation with the Miocene sequence of the nearby basin of Chiang Muan, the large-bodied hominoid ( Khoratpithecus chiangmuanensis )–bearing levels can be precisely dated between 12.4 and 12.2 Ma.

First Hominoid from the Late Miocene of the Irrawaddy Formation (Myanmar)

For over a century, a Neogene fossil mammal fauna has been known in the Irrawaddy Formation in central Myanmar. Unfortunately, the lack of accurately located fossiliferous sites and the absence of hominoid fossils have impeded paleontological studies. Here we describe the first hominoid found in Myanmar together with a Hipparion (s.l.) associated mammal fauna from Irrawaddy Formation deposits dated between 10.4 and 8.8 Ma by biochronology and magnetostratigraphy. This hominoid documents a new species of Khoratpithecus, increasing thereby the Miocene diversity of southern Asian hominoids. The composition of the associated fauna as well as stable isotope data on Hipparion (s.l.) indicate that it inhabited an evergreen forest in a C3-plant environment. Our results enlighten that late Miocene hominoids were more regionally diversified than other large mammals, pointing towards regionally-bounded evolution of the representatives of this group in Southeast Asia. The Irrawaddy Formation, with its extensive outcrops and long temporal range, has a great potential for improving our knowledge of hominoid evolution in Asia.

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.

Chronology of the Eocene continental deposits of Africa: Magnetostratigraphy and biostratigraphy of the El Kohol and Glib Zegdou Formations, Algeria

Despite numerous discoveries that have considerably enriched the African-Arabian Tertiary fossil record over the last decades, our knowledge of the evolutionary history of many continental African vertebrate groups during the Paleogene period remains inadequate, particularly when it is compared with the fossil records of Europe or North America. The Eocene Epoch in Africa is especially poorly documented, being restricted to few fossiliferous localities. Our understanding of the early Tertiary emergence, diversification, and paleobiogeographic history of African-Arabian mammals has been further hindered by the lack of a precise temporal framework for these sites. We conducted magnetostratigraphic analyses, associated with biostratigraphic studies, in the fossiliferous sequences exposed in the northwestern Hammadas of the Saharan Platform in the Glib Zegdou area and in the Saharan Atlas at the El Kohol locality (Algeria) to further define the age of these Eocene continental deposits. Based on biostratigraphic constraints, the six polarity zones identified in the El Kohol section can be correlated with chrons C24n to C22r, providing the first direct age estimates for the El Kohol fossiliferous strata between 52 and 51 Ma. Correlation to the geomagnetic polarity time scale, using previously published biostratigraphic data for the Glib Zegdou fauna, suggests an age ranging between 49 and 45 Ma for this section. The high-resolution magnetostratigraphic study of the poorly known continental Eocene Epoch of Algeria provides new insights into the early Tertiary stratigraphy of northwest Africa. The placement of the Algerian localities into a consistent chronological framework constitutes considerable advancement to achieve biostratigraphic correlation of the Paleogene African-Arabian mammal localities.

First record of a chalicothere from the Miocene of Myanmar

Here we describe the first record of a chalicothere from the Miocene of Myanmar. The chalicothere, documented by a partial mandible, was unearthed from the lower portion of the Irrawaddy Formation in the region of Magway, Central Myanmar. The Burmese material belongs to an early late Miocene fauna which recently yielded hominoid remains attributed to Khoratpithecus. The specimen, which is attributed to a chalicotheriine, does not reliably match with any described Miocene Eurasian species of this subfamily, suggesting the possibility it belongs to a new taxon. The discovery of a chalicotheriine in the surroundings of Magway contributes to the hypothesis that closed habitats were an important component of the paleoenvironment of Khoratpithecus.

New fossils from the Paleogene of central Libya illuminate the evolutionary history of endemic African anomaluroid rodents

Anomaluroid rodents show interesting biogeographic and macroevolutionary patterns, although their fossil record is meager and knowledge of the natural history of extant members of the clade remains inadequate. Living anomaluroids (Anomaluridae) are confined to equatorial parts of western and central Africa, but the oldest known fossil anomaluroid (Pondaungimys) comes from the late middle Eocene of Myanmar. The first appearance of anomaluroids in the African fossil record coincides with the first appearances of hystricognathous rodents and anthropoid primates there. Both of the latter taxa are widely acknowledged to have originated in Asia, suggesting that anomaluroids may show a concordant biogeographic pattern. Here we describe two new taxa of African Paleogene anomaluroids from sites in the Sirt Basin of central Libya. These include a new Eocene species of the nementchamyid genus Kabirmys, which ranks among the oldest African anomaluroids recovered to date, and a new genus and species of Anomaluridae from the early Oligocene, which appears to be closely related to extant Zenkerella, the only living non-volant anomalurid. Phylogenetic analyses incorporating the new Libyan fossils suggest that anomaluroids are not specially related to Zegdoumyidae, which are the only African rodents known to antedate the first appearance of anomaluroids there. The evolution of gliding locomotion in Anomaluridae appears to conflict with traditional assessments of relationships among living anomalurid taxa. If the historically accepted division of Anomaluridae into Anomalurinae (extant and Miocene Anomalurus and Miocene Paranomalurus) and Zenkerellinae (extant and Miocene Zenkerella and extant Idiurus) is correct, then either gliding locomotion evolved independently in Anomalurinae and Idiurus or non-volant Zenkerella evolved from a gliding ancestor. Anatomical data related to gliding in Anomaluridae are more consistent with a nontraditional systematic arrangement, whereby non-volant Zenkerella is the sister group of a clade including both Anomalurus and Idiurus.

Uniquely derived upper molar morphology of Eocene Amphipithecidae (Primates: Anthropoidea): Homology and phylogeny

The extinct Southeast Asian primate family Amphipithecidae is regularly cited in discussions of anthropoid origins, but its phylogenetic position remains controversial. In part, the lack of consensus regarding amphipithecid relationships can be attributed to uncertainty regarding the homology of upper molar structures in this group. Here, we describe a virtually pristine upper molar of Pondaungia cotteri from the late middle Eocene Pondaung Formation of Myanmar, which is the first example of a relatively unworn and well-preserved amphipithecid upper molar ever recovered. The distolingual upper molar cusp in this new specimen of Pondaungia appears to be a lingually displaced and enlarged metaconule, rather than a hypocone or pseudohypocone as previous workers have thought. Reassessment of the upper molar morphology of other amphipithecids and putative amphipithecids reveals a very similar pattern in Siamopithecus, Myanmarpithecus and Ganlea, all of which are interpreted as having upper molars showing many of the same derived features apparent in Pondaungia. In contrast, the upper molar morphology of Bugtipithecus diverges radically from that of undoubted amphipithecids, and the latter taxon is excluded from Amphipithecidae on this basis. Phylogenetic analyses of several character-taxon matrices culled from the recent literature and updated to reflect the new information on amphipithecid upper molar morphology yield similar results. Consensus tree topologies derived from these analyses support amphipithecid monophyly and stable relationships within Amphipithecidae. Amphipithecids appear to be stem members of the anthropoid clade.

A stem bat (Chiroptera: Palaeochiropterygidae) from the late middle Eocene of northern Anatolia: implications for the dispersal and palaeobiology of early bats

Fragmentary remains of an Eocene bat are described from the middle Eocene Lülük Member of the Uzunçarşidere Formation on the Pontide terrane, in what is now north-central Anatolia. The new taxon most closely resembles the palaeochiropterygids Lapichiropteryx and Stehlinia in terms of its known dental morphology, and it is referred to the stem chiropteran family Palaeochiropterygidae on this basis. Geological and palaeontological data indicate that the Pontide terrane was an island situated along the northern margin of Neotethys during the middle Eocene. The presence of a late-surviving stem chiropteran in an island context potentially illuminates dispersal patterns and capabilities among the earliest bats, which already enjoyed a nearly global distribution by the early Eocene. Other palaeochiropterygids for which postcranial material is known share little in common with extant bats that are capable of long-range dispersal across open water. The new Turkish bat taxon is consistent with a hypothetical dispersal corridor between Western Europe and India via islands on the northern margin of Neotethys and suggests a larger range of skeletal and locomotor variation within Palaeochiropterygidae than is currently recognised.