Pieter Missiaen

Early Eocene primates from Gujarat, India.

The oldest euprimates known from India come from the Early Eocene Cambay Formation at Vastan Mine in Gujarat. An Ypresian (early Cuisian) age of approximately 53Ma (based on foraminifera) indicates that these primates were roughly contemporary with, or perhaps predated, the India-Asia collision. Here we present new euprimate fossils from Vastan Mine, including teeth, jaws, and referred postcrania of the adapoids Marcgodinotius indicus and Asiadapis cambayensis. They are placed in the new subfamily Asiadapinae (family Notharctidae), which is most similar to primitive European Cercamoniinae such as Donrussellia and Protoadapis. Asiadapines were small primates in the size range of extant smaller bushbabies. Despite their generally very plesiomorphic morphology, asiadapines also share a few derived dental traits with sivaladapids, suggesting a possible relationship to these endemic Asian adapoids. In addition to the adapoids, a new species of the omomyid Vastanomys is described. Euprimate postcrania described include humeri, radii, femora, calcanei, and tali, most of which show typical notharctid features and are probably attributable to asiadapines. Anatomical features of the limb elements indicate that they represent active arboreal quadrupedal primates. At least one calcaneus is proximally shorter and distally longer than the others, resembling eosimiids in this regard, a relationship that, if confirmed, would also suggest an Asian or southeast Asian faunal connection. Isolated teeth from Vastan Mine recently attributed to a new eosimiid, Anthrasimias gujaratensis, appear to provide that confirmation. However, their attribution to Eosimiidae is equivocal. They are similar to teeth here tentatively referred to Marcgodinotius, hence A. gujaratensis may be a junior synonym of M. indicus. Corroboration of eosimiids at Vastan requires more compelling evidence. Although definitive conclusions are premature, available evidence suggests that the Vastan adapoids, at least, were derived from western European stock that reached India near the Paleocene-Eocene boundary.

High bat (Chiroptera) diversity in the Early Eocene of India

The geographic origin of bats is still unknown, and fossils of earliest bats are rare and poorly diversified, with, maybe, the exception of Europe. The earliest bats are recorded from the Early Eocene of North America, Europe, North Africa and Australia where they seem to appear suddenly and simultaneously. Until now, the oldest record in Asia was from the Middle Eocene. In this paper, we report the discovery of the oldest bat fauna of Asia dating from the Early Eocene of the Cambay Formation at Vastan Lignite Mine in Western India. The fossil taxa are described on the basis of well-preserved fragments of dentaries and lower teeth. The fauna is highly diversified and is represented by seven species belonging to seven genera and at least four families. Two genera and five species are new. Three species exhibit very primitive dental characters, whereas four others indicate more advanced states. Unexpectedly, this fauna presents strong affinities with the European faunas from the French Paris Basin and the German Messel locality. This could result from the limited fossil record of bats in Asia, but could also suggest new palaeobiogeographic scenarios involving the relative position of India during the Early Eocene.

Early Eocene lagomorph (Mammalia) from Western India and the early diversification of Lagomorpha

We report the oldest known record of Lagomorpha, based on distinctive, small ankle bones (calcaneus and talus) from Early Eocene deposits (Middle Ypresian equivalent, ca 53 Myr ago) of Gujarat, west-central India. The fossils predate the oldest previously known crown lagomorphs by several million years and extend the record of lagomorphs on the Indian subcontinent by 35 Myr. The bones show a mosaic of derived cursorial adaptations found in gracile Leporidae (rabbits and hares) and primitive traits characteristic of extant Ochotonidae (pikas) and more robust leporids. Together with gracile and robust calcanei from the Middle Eocene of Shanghuang, China, also reported here, the Indian fossils suggest that diversification within crown Lagomorpha and possibly divergence of the family Leporidae were already underway in the Early Eocene.

Early Eocene (Ypresian) continental vertebrate assemblage from India, with description of a new anthracobunid (Mammalia, Tethytheria)

K. D. ROSE, T. SMITH, R. S. RANA, A. SAHNI, H. SINGH, P. MISSIAEN, and A. FOLIE Center for Functional Anatomy and Evolution, Johns Hopkins University School of Medicine, Baltimore, Maryland 21205, kdrose@jhmi.edu; Department of Paleontology, Royal Belgian Institute of Natural Sciences, B-1000 Brussels, Belgium, Thierry.Smith@naturalsciences.be, Annelise.Folie@naturalsciences.be; Department of Geology, H.N.B. Garhwal University, Srinagar 246175, Uttaranchal, India, rs_rana_hnbgu@yahoo.com; Centre of Advanced Study in Geology, Panjab University, Chandigarh-160014, India, ashok.sahni@gmail.com; Research Unit of Paleontology, University of Ghent, B-9000 Ghent, Belgium, Pieter.Missiaen@ugent.be

Early Eocene fossils suggest that the mammalian order Perissodactyla originated in India

Cambaytheres (Cambaytherium, Nakusia and Kalitherium) are recently discovered early Eocene placental mammals from the Indo-Pakistan region. They have been assigned to either Perissodactyla (the clade including horses, tapirs and rhinos, which is a member of the superorder Laurasiatheria) or Anthracobunidae, an obscure family that has been variously considered artiodactyls or perissodactyls, but most recently placed at the base of Proboscidea or of Tethytheria (Proboscidea+Sirenia, superorder Afrotheria). Here we report new dental, cranial and postcranial fossils of Cambaytherium, from the Cambay Shale Formation, Gujarat, India (~54.5 Myr). These fossils demonstrate that cambaytheres occupy a pivotal position as the sister taxon of Perissodactyla, thereby providing insight on the phylogenetic and biogeographic origin of Perissodactyla. The presence of the sister group of perissodactyls in western India near or before the time of collision suggests that Perissodactyla may have originated on the Indian Plate during its final drift toward Asia.

First skull of Orthaspidotherium edwardsi (Mammalia, “Condylarthra”) from the late Paleocene of Berru (France) and phylogenetic affinities of the enigmatic European family Pleuraspidotheriidae

ABSTRACT Among the archaic ‘ungulates,’ pleuraspidotheriids are well documented by skulls and postcranial elements of Pleuraspidotherium from Berru and Cernay-lès-Reims (late Paleocene, Paris Basin, France). Nevertheless, the relationships of pleuraspidotheriids (i.e., Pleuraspidotherium, Orthaspidotherium, and Hilalia) to other ‘condylarths’ have not been conclusively settled. They have been related to the typically North American Meniscotheriidae, Phenacodontidae, and Mioclaenidae, and even to the modern ungulates Perissodactyla and Artiodactyla. We here provide additional data, especially from the first complete skull of Orthaspidotherium, and highlight a mosaic of plesiomorphic and derived characters that distinguish Pleuraspidotheriidae from all other ‘ungulates.’ Their basicranial morphology approximates the ancestral morphotype of artiodactyls, but this results from symplesiomorphies. Pleuraspidotheriids exhibit the following synapomorphies: strong processes on the petrosal and enclosure of the facial nerve by the tympanic process and tympanohyal. Although their dentition resembles that of perissodactyls, meniscotheriids, and phenacodontids, the development of a pseudohypocone from the metaconule unambiguously shows that these similarities arose by convergence. Other evolutionary tendencies are the strong reduction of the hypoconulid, molarization of last premolars, development of lingual cingulum on M2, lengthening of the snout, and presence of diastemata. The pleuraspidotheriid tarsal morphology is primitive and not uncommon among ‘condylarths’ but presents some synapomorphies such as a calcaneum with a transverse cuboid facet and an astragalus with a squatting facet, cotylar fossa, and sustentacular hinge. Our study does not indicate a close relationship of Pleuraspidotheriidae with any of the archaic or modern ‘ungulates’ mentioned above. Instead, primitive arctocyonids could represent a possible ancestral morphotype for pleuraspidotheriids.

The Gashatan (Late Paleocene) Mammal Fauna from Subeng, Inner Mongolia, China

The Paleocene—Eocene boundary is of particular importance for the evolution of mammals and the poorly known Asian mammal faunas from this period have received much attention. The late Paleocene Subeng site in Inner Mongolia (China) has come under study only recently, and here we present the first complete description of its mammal fauna. Two new species are described, the neoplagiaulacid multituberculate Mesodmops tenuis sp. nov. and the praolestine nyctitheriid Bumbanius ningi sp. nov., representing stratigraphic range extensions of the respective genera into the Paleocene. Previously unknown parts of the dentition are described here for the eurymylid Eomylus bayanulanensis, the sarcodontid Hyracolestes ermineus, the cimolestid Tsaganius ambiguus, the carpolestid Subengius mengi, as well as the femur of the mesonychid Dissacus serratus. For most taxa, the new specimens from Subeng provide new phylogenetic and/or biostratigraphic information. We confirm the inclusion of Hyracolestes in the Sarcodontinae and elevate this group to the rank of family, the Sarcodontidae, separate from Micropternodontidae. In the case of Subengius mengi an updated cladistic analysis of carpolestids supports the hypothesis that Subengius is derived from an evolved Elphidotarsius-like ancestor in the early to middle Tiffanian of North America. A total of 17 species is identified, including well-known biostratigraphic markers for the late Paleocene Gashatan Asian Land Mammal Age such as Lambdopsalis bulla, Prionessus sp., Palaeostylops iturus, Pseudictops lophiodon, Tribosphenomys minutus, and Dissacus serratus. We propose that the Gashatan faunas are less endemic than previously thought, and result from a significant exchange with North American faunas from the late Paleocene.

Asian gliriform origin for arctostylopid mammals.

Arctostylopids are common in Late Palaeocene and Early Eocene mammal faunas of Asia, but they are rare in North America. From the time of their first discovery, arctostylopids were grouped with the strictly endemic South American Notoungulata based on their strikingly similar dental morphology. This relationship was challenged based on dental morphology of more recently discovered arctostylopids and on supposed tarsal morphology (based on unassociated material) of the Asian arctostylopid Palaeostylops. Therefore, Arctostylopidae were placed in a separate order, Arctostylopida, of enigmatic affinities. Many authors, however, continue to unite Arctostylopidae and Notoungulata based on dental similarities. In the Late Palaeocene Subeng site in Inner Mongolia, we identified the Palaeostylops tarsals based on their size and abundance. This identification is supported by comparison to unpublished Arctostylops tarsals from the North American Clarkforkian, derived from a semi-articulated skeleton also including dental material. Tarsal morphology shows moderate similarity to the gliriform Pseudictops, and strong resemblance to the tarsally conservative gliroid Rhombomylus. Hence, Arctostylopidae may best be grouped with Asian non-gliroid Gliriformes, which we interpret as having dispersed into North America in the Late Palaeocene.

Craniodental and postcranial morphology of Indohyaenodon raoi from the early eocene of india, and its implications for ecology, phylogeny, and biogeography of hyaenodontid mammals

ABSTRACT New remains of the early Eocene hyaenodontid Indohyaenodon raoi are described from the Vastan Lignite Mine in Gujarat, western India, including the first known rostrum, upper dentition, and postcrania, substantially expanding our knowledge of the species and providing insights into its functional morphology and relationships. Craniodental morphology suggests that I. raoi had a broad diet, including non-vertebrate material as well as flesh of a diversity of prey species. Postcranial morphology is broadly similar to that of other early hyaenodontids and suggests a scansorial locomotor repertoire. Dental morphology indicates that I. raoi is closely related to other South Asian hyaenodontids, with shared features including strong cingula, narrow premolars, and a reduced P4 protocone. We present the most comprehensive phylogenetic analysis of Hyaenodontidae to date, which corroborates this relationship but finds South Asian hyaenodontids to be the stem of a group that includes most African hyaenodontids. This and other higher-level relationships within Hyaenodontidae are, however, weakly supported, and substantially different alternative hypotheses of relationships are not significantly less parsimonious, reflecting strong character conflict. Factors contributing to this conflict include the isolation of hyaenodontid faunas on different continents during much of the Eocene, canalization and simplification of carnivorous dentitions, and a lack of non-dental material for critical hyaenodontid groups. The new phylogeny is consistent with either an African or an Asian origin for the group.

New Early Eocene Tapiromorph Perissodactyls from the Ghazij Formation of Pakistan, with Implications for Mammalian Biochronology in Asia

Early Eocene mammals from Indo-Pakistan have only recently come under study. Here we describe the first tapiromorph perissodactyls from the subcontinent. Gandheralophus minor gen. et sp. nov. and G. robustus sp. nov. are two species of Isectolophidae differing in size and in reduction of the anterior dentition. Gandheralophus is probably derived from a primitive isectolophid such as Orientolophus hengdongensis from the earliest Eocene of China, and may be part of a South Asian lineage that also contains Karagalax from the middle Eocene of Pakistan. Two specimens are referred to a new, unnamed species of Lophialetidae. Finally, a highly diagnostic M3 and a molar fragment are described as the new eomoropid chalicothere Litolophus ghazijensis sp. nov. The perissodactyls described here, in contrast to most other mammalian groups published from the early Eocene of Indo-Pakistan, are most closely related to forms known from East and Central Asia, where Eocene tapiromorphs are diverse and biochronologically important. Our results therefore allow the first biochronological correlation between early Eocene mammal faunas in Indo-Pakistan and the rest of Asia. We suggest that the upper Ghazij Formation of Pakistan is best correlated with the middle or late part of the Bumbanian Asian Land Mammal Age, while the Kuldana and Subathu Formations of Pakistan and India are best correlated with the Arshantan Asian Land Mammal Age.