A new primate from the late Eocene of Vietnam illuminates unexpected strepsirrhine diversity and evolution in Southeast Asia.

Sivaladapidae is a poorly known Asian strepsirrhine family originally discovered in Miocene sediments of the Indian subcontinent. Subsequent research has considerably increased the diversity, temporal range, and geographical distribution of this group, now documented from China, Thailand, Myanmar, Pakistan, and India and whose earliest representatives date back to the Middle Eocene. We present here a new taxon of sivaladapid from the Na Duong coal mine in the Latest Middle Eocene-Late Eocene of Vietnam. It represents the first Eocene primate from Vietnam and the first medium-sized mammal recovered from this locality, thus documenting a completely new part of the Na Duong paleobiodiversity. This taxon is the largest sivaladapid ever found with an estimated body weight of 5.3 kg and it represents a new subfamily of sivaladapids in exhibiting a very peculiar combination of dental features yet unknown in the fossil record of the family (e.g., retention of four premolars, high-crowned molars with accentuated bunodonty and extreme crest reduction). Besides documenting a complete new part of sivaladapid evolution, its primitive dental formula and derived features shared with the Early Eocene Asiadapidae reinforce the hypothesis of a basal branching of sivaladapids among strepsirrhines.

Endocasts and brain evolution in Anthracotheriidae (Artiodactyla, Hippopotamoidea).

Anthracotheres are a fossil family of 'Suiformes' from the Old World, North and Central America. They are known from the middle Eocene to the late Pliocene, and are suggested to be the stem group of Hippopotamidae. Yet, their soft anatomy remains poorly known. In this study we describe the virtual endocast of the late Oligocene anthracothere Microbunodon minimum, reconstructed using microtomography, as well as the natural endocast of Merycopotamus medioximus from the late Miocene. These are the first anthracothere endocasts ever described. Particular attention is given to the relative proportions of the brain, the neocortex, the cerebellum and the olfactory bulbs. The 'backward shift' of the pituitary of M. minimum, and the possible presence of a K lobe in M. medioximus, is discussed. Previous statements that some endocranial characters were subject to convergence among mammals are also corroborated.

Hippos stem from the longest sequence of terrestrial cetartiodactyl evolution in Africa.

According to molecular data, hippopotamuses and cetaceans form a clade excluding other extant cetartiodactyls. Despite a wealth of spectacular specimens documenting cetacean evolution, this relationship remains poorly substantiated by the fossil record. Indeed, the evolutionary path leading from the hippo-cetacean ancestor to Hippopotamidae is plagued by missing fossil data and phylogenetic uncertainties. Only an origination within the extinct anthracotheres is compatible with molecular results, substantial filling of phyletic gaps and recent discoveries of early Miocene hippopotamids. Yet, the anthracothere stock that gave rise to Hippopotamidae has not been identified. Consequently, recent phylogenetic accounts do not properly integrate the anthracotheriid hypothesis, and relate Hippopotamidae to a stretched ghost lineage and/or close to Suina. Here we describe a new anthracothere from Lokone (Kenya) that unambiguously roots the Hippopotamidae into a well-identified group of bothriodontines, the first large mammals to invade Africa. The hippos are deeply anchored into the African Paleogene.

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.

Covariation between midline cranial base, lateral basicranium, and face in modern humans and chimpanzees: a 3D geometric morphometric analysis.

Previous studies showed that in modern humans the basicranium is formed of two modules: the midline cranial base and the lateral basicranium which are integrated with the face in very different ways. The study of the relationship between these structures is of prime interest in the context of hominids craniofacial evolutionary history. In this study, we aim to test if the relationship between the midline cranial base and the face on one hand and the lateral basicranium and the face on the other hand are qualitatively and quantitatively different in modern humans and chimpanzees: two phylogenetically close but morphologically different hominids. This work is performed using three-dimensional (3D) landmarks to take into account the face and basicranium 3D shape. Modern humans and chimpanzees both exhibit a significant relationship between lateral basicranium and face, and a nonsignificant relationship between midline cranial base and face. However, the patterns of integration are different for the two species. These results underscore the essential role of the lateral basicranial shape in the setting of the facial morphology in modern humans and chimpanzees. The important differences in the patterns of integration may be related to the genetic, developmental, and functional requirements of each taxon, acquired along their respective evolution. From a common, tight, relationship between lateral basicranium, and face, each taxon may develop different patterns of integration in order to adapt to particular functions and morphologies.

First record of a parapithecid primate from the Oligocene of Kenya.

Recent excavations in northwestern Kenya have recovered a vertebrate fauna of late early or early late Oligocene age. Among the mammal remains, a fragmentary lower jaw and an isolated upper molar have been attributed to a small primate, Lokonepithecus manai gen. et sp. nov. Lokonepithecus is a primitive member of the Parapithecidae and possibly most closely related to Apidium from the Fayum. The new primate from Kenya is the youngest parapithecid known and its occurrence in the Oligocene of Kenya suggests that sub-Saharan Africa probably played a major role in the evolutionary history of several groups of mammals.

Eocene bunoselenodont Artiodactyla from southern Thailand and the early evolution of Ruminantia in South Asia.

Although Asia is thought to have played a critical role in the basal radiation of Ruminantia, the fossil record of early selenodont artiodactyls remains poorly documented in this region. Dental remains of a new bunoselenodont artiodactyl are described from the late Eocene of Krabi, southern Thailand. This new form, Krabitherium waileki gen. et sp. nov, is tentatively referred to the Tragulidae (Ruminantia) on the basis of several dental features, including a weak Tragulus fold and the presence of a deep groove on the anterior face of the entoconid. Although this new form is suggestive of the enigmatic? Gelocus gajensis Pilgrim 1912 from the "base of the Gaj" (lower Chitarwata Formation) of the Bugti Hills (Central Pakistan), K. waileki most likely represents an early representative of a relatively bunodont group of tragulids that includes the genus Dorcabune, known from the Miocene of south Asia. This addition to the Eocene record of early ruminants attests to the antiquity of the group in Southeast Asia and lends support to the hypothesis that the Tragulidae represents one of the first offshoots in the evolutionary history of Ruminantia.

New primates from the late Eocene of Thailand: a contribution to primate diversity in the Paleogene of Asia.

We report the discovery of two new primates from the late Eocene Krabi Basin, southern Thailand. One isolated upper molar displays morphological features (protocone united with hypocone by the prehypocrista, postprotocrista extended distobuccally) suggesting possible phylogenetic relationships with Amphipithecidae, while an isolated lower molar is tentatively referred to as a new tarsiiform, mainly on the basis of its paraconid and entoconid morphology. Although very scarce, these remains expand the record of Paleogene primates in Southeast Asia, and testify to their successful radiation in that area.

The anthropoid status of a primate from the late middle Eocene Pondaung Formation (Central Myanmar): tarsal evidence.

Primate dental and postcranial remains from the Eocene Pondaung Formation (Myanmar) have been the subject of considerable confusion since their initial discoveries, and their anthropoid status has been widely debated. We report here a well preserved primate talus discovered in the Segyauk locality near Mogaung that displays derived anatomical features typical of haplorhines, notably anthropoids, and lacks strepsirhine synapomorphies. Linear discriminant and parsimony analyses indicate that the talus from Myanmar is more similar structurally to those of living and extinct anthropoids than to those of adapiforms, and its overall osteological characteristics further point to arboreal quadrupedalism. Regressions of talar dimensions versus body mass in living primates indicate that this foot bone might have belonged to Amphipithecus. This evidence supports hypotheses favoring anthropoid affinities for the large-bodied primates from Pondaung and runs contrary to the hypothesis that Pondaungia and Amphipithecus are strepsirhine adapiforms.

Oligocene sivaladapid primate from the Bugti Hills (Balochistan, Pakistan) bridges the gap between Eocene and Miocene adapiform communities in Southern Asia.

A new species of Guangxilemur (Sivaladapidae, Adapiformes) is described from the early Oligocene Chitarwata Formation (Bugti Member) of the Bugti Hills, Sulaiman geological Province, Balochistan, Pakistan. Guangxilemur singsilai n. sp. provides further diagnostic morphological characters from its newly described upper and lower dentitions, confirming its intermediate phylogenetic position between Eocene and Miocene Asian sivaladapid adapiforms. G. singsilai possesses moderately developed shearing and puncturing molar features and maintains lingual cusps on upper molars as in Eocene hoanghoniines; in contrast, it possesses a typical molariform P(4) as in Miocene sivaladapines. The important paleogeographic changes that have affected South Asia during the Tertiary (related to the collision between the Indian and Eurasian Plates) have played a critical role in reforming circulation and climatic differentiation. The presence in Pakistan of an unique and well-diversified Oligocene primate fauna, clearly demonstrates that South Asia maintained favourable environmental conditions during the middle Caenozoic global climatic deterioration that coincides with drastic changes in faunal structure on the whole Holarctic Province, including the extinction of adapiform primates.