A morphometric analysis of early Eocene Euprimate tarsals from Gujarat, India.

Early Eocene primate postcranial bones from the Vastan lignite mine of Gujarat, India, have proven useful for understanding the haplorhine and strepsirrhine divergence. Previous analyses of material assigned to Asiadapidae supported interpretations that these primates were generalized arboreal quadrupeds, while the omomyid Vastanomys was likely to have been more proficient leaper than asiadapids. More recent examinations of long bone cross-sectional properties and calcaneal elongation have complicated the behavioral interpretations of these fossils. This study examines whole talar and calcaneal morphology of the Vastan material to refine the locomotor reconstructions of these fossils. A comparative sample of extant primate species representing various locomotor behaviors was obtained by accessing surface models from MorphoSource.org. Surface models of fossil specimens attributed to Asiadapis cambayensis, Marcgodinotius indicus, and Vastanomys major were generated from micro-computed tomography scans. A morphological analysis was carried out using weighted spherical harmonics, a Fourier-based method that represents surfaces using coefficients associated with a common set of spherical harmonic functions. The coefficients describing each surface were then used as shape variables in a principal components analysis. Significant differences between locomotor groups were assessed using nonparametric tests. Results from extant comparative samples show that locomotor behavior can be predicted from both talar and calcaneal morphology when phylogenetic relationships are known. Consistent with previous analyses, our results indicate that Asiadapis cambayensis and Marcgodinotius indicus were likely arboreal quadrupeds with some leaping capabilities. Vastanomys major is reconstructed as an arboreal quadruped with greater leaping proficiency than its asiadapid counterparts based on its talar morphology.

Cladogenesis and replacement in the fossil record of Microsyopidae (?Primates) from the southern Bighorn Basin, Wyoming.

The early Eocene of the southern Bighorn Basin, Wyoming, is notable for its nearly continuous record of mammalian fossils. Microsyopinae (?Primates) is one of several lineages that shows evidence of evolutionary change associated with an interval referred to as Biohorizon A. Arctodontomys wilsoni is replaced by a larger species, Arctodontomys nuptus, during the biohorizon interval in what is likely an immigration/emigration or immigration/local extinction event. The latter is then superseded by Microsyops angustidens after the end of the Biohorizon A interval. Although this pattern has been understood for some time, denser sampling has led to the identification of a specimen intermediate in morphology between A. nuptus and M. angustidens, located stratigraphically as the latter is appearing. Because specimens of A. nuptus have been recovered approximately 60 m above the appearance of M. angustidens, it is clear that A. nuptus did not suffer pseudoextinction. Instead, evidence suggests that M. angustidens branched off from a population of A. nuptus, but the latter species persisted. This represents possible evidence of cladogenesis, which has rarely been directly documented in the fossil record. The improved understanding of both evolutionary transitions with better sampling highlights the problem of interpreting gaps in the fossil record as punctuations.

New fossils from Tadkeshwar Mine (Gujarat, India) increase primate diversity from the early Eocene Cambay Shale.

Several new fossil specimens from the Cambay Shale Formation at Tadkeshwar Lignite Mine in Gujarat document the presence of two previously unknown early Eocene primate species from India. A new species of Asiadapis is named based on a jaw fragment preserving premolars similar in morphology to those of A. cambayensis but substantially larger. Also described is an exceptionally preserved edentulous dentary (designated cf. Asiadapis, unnamed sp. nov.) that is slightly larger and much more robust than previously known Cambay Shale primates. Its anatomy most closely resembles that of Eocene adapoids, and the dental formula is the same as in A. cambayensis. A femur and calcaneus are tentatively allocated to the same taxon. Although the dentition is unknown, exquisite preservation of the dentary of cf. Asiadapis sp. nov. enables an assessment of masticatory musculature, function, and gape adaptations, as well as comparison with an equally well-preserved dentary of the asiadapid Marcgodinotius indicus, also from Tadkeshwar. The new M. indicus specimen shows significant gape adaptations but was probably capable of only weak bite force, whereas cf. Asiadapis sp. nov. probably used relatively smaller gapes but could generate relatively greater bite forces.

New euprimate postcrania from the early Eocene of Gujarat, India, and the strepsirrhine-haplorhine divergence.

The oldest primates of modern aspect (euprimates) appear abruptly on the Holarctic continents during a brief episode of global warming known as the Paleocene-Eocene Thermal Maximum, at the beginning of the Eocene (∼56 Ma). When they first appear in the fossil record, they are already divided into two distinct clades, Adapoidea (basal members of Strepsirrhini, which includes extant lemurs, lorises, and bushbabies) and Omomyidae (basal Haplorhini, which comprises living tarsiers, monkeys, and apes). Both groups have recently been discovered in the early Eocene Cambay Shale Formation of Vastan lignite mine, Gujarat, India, where they are known mainly from teeth and jaws. The Vastan fossils are dated at ∼54.5 Myr based on associated dinoflagellates and isotope stratigraphy. Here, we describe new, exquisitely preserved limb bones of these Indian primates that reveal more primitive postcranial characteristics than have been previously documented for either clade, and differences between them are so minor that in many cases we cannot be certain to which group they belong. Nevertheless, the small distinctions observed in some elements foreshadow postcranial traits that distinguish the groups by the middle Eocene, suggesting that the Vastan primates-though slightly younger than the oldest known euprimates-may represent the most primitive known remnants of the divergence between the two great primate clades.

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.

Body size and premolar evolution in the early-middle eocene euprimates of Wyoming.

The earliest euprimates to arrive in North America were larger-bodied notharctids and smaller-bodied omomyids. Through the Eocene, notharctids generally continued to increase in body size, whereas omomyids generally radiated within small- and increasingly mid-sized niches in the middle Eocene. This study examines the influence of changing body size and diet on the evolution of the lower fourth premolar in Eocene euprimates. The P4 displays considerable morphological variability in these taxa. Despite the fact that most studies of primate dental morphology have focused on the molars, P4 can also provide important paleoecological insights. We analyzed the P4 from 177 euprimate specimens, representing 35 species (11 notharctids and 24 omomyids), in three time bins of approximately equal duration: early Wasatchian, late Wasatchian, and Bridgerian. Two-dimensional surface landmarks were collected from lingual photographs, capturing important variation in cusp position and tooth shape. Disparity metrics were calculated and compared for the three time bins. In the early Eocene, notharctids have a more molarized P4 than omomyids. During the Bridgerian, expanding body size range of omomyids was accompanied by a significant increase in P4 disparity and convergent evolution of the semimolariform condition in the largest omomyines. P4 morphology relates to diet in early euprimates, although patterns vary between families.

New fossils of the oldest North American euprimate Teilhardina brandti (Omomyidae) from the paleocene-eocene thermal maximum.

More than 25 new specimens of Teilhardina brandti, one of the oldest known euprimates, are reported from earliest Eocene strata of the southern Bighorn Basin, Wyoming. The new fossils include the first upper dentitions, a dentary showing the lower dental formula for the first time, and the first postcrania ascribed to T. brandti (tarsals and terminal phalanges). The elongated navicular and long talar neck suggest that T. brandti was an active arboreal quadruped, and the terminal phalanges constitute the oldest evidence for nails in Euprimates. Phylogenetic analysis incorporating the new data indicates that T. brandti is more derived than T. belgica but less so than T. americana. The hypothesis that Teilhardina originated in Asia (T. asiatica) and dispersed westward to Europe (T. belgica) and then to North America (T. brandti and T. magnoliana) during the earliest Eocene Paleocene-Eocene Thermal Maximum is most consistent with available evidence, including the relative age of fossil samples and their stage of evolution.

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.

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.

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.

Rapid Asia-Europe-North America geographic dispersal of earliest Eocene primate Teilhardina during the Paleocene-Eocene Thermal Maximum.

True primates appeared suddenly on all three northern continents during the 100,000-yr-duration Paleocene-Eocene Thermal Maximum at the beginning of the Eocene, approximately 55.5 mya. The simultaneous or nearly simultaneous appearance of euprimates on northern continents has been difficult to understand because the source area, immediate ancestors, and dispersal routes were all unknown. Now, omomyid haplorhine Teilhardina is known on all three continents in association with the carbon isotope excursion marking the Paleocene-Eocene Thermal Maximum. Relative position within the carbon isotope excursion indicates that Asian Teilhardina asiatica is oldest, European Teilhardina belgica is younger, and North American Teilhardina brandti and Teilhardina americana are, successively, youngest. Analysis of morphological characteristics of all four species supports an Asian origin and a westward Asia-to-Europe-to-North America dispersal for Teilhardina. High-resolution isotope stratigraphy indicates that this dispersal happened in an interval of approximately 25,000 yr. Rapid geographic dispersal and morphological character evolution in Teilhardina reported here are consistent with rates observed in other contexts.

Affinities of 'hyopsodontids' to elephant shrews and a Holarctic origin of Afrotheria.

Macroscelideans (elephant shrews or sengis) are small-bodied (25-540 g), cursorial (running) and saltatorial (jumping), insectivorous and omnivorous placental mammals represented by at least 15 extant African species classified in four genera. Macroscelidea is one of several morphologically diverse but predominantly African placental orders classified in the superorder Afrotheria by molecular phylogeneticists. The distribution of modern afrotheres, in combination with a basal position for Afrotheria within Placentalia and molecular divergence-time estimates, has been used to link placental diversification with the mid-Cretaceous separation of South America and Africa. Morphological phylogenetic analyses do not support Afrotheria and the fossil record favours a northern origin of Placentalia. Here we describe fossil postcrania that provide evidence for a close relationship between North American Palaeocene-Eocene apheliscine 'hyopsodontid' 'condylarths' (early ungulates or hoofed mammals) and extant Macroscelidea. Apheliscine postcranial morphology is consistent with a relationship to other ungulate-like afrotheres (Hyracoidea, Proboscidea) but does not provide support for a monophyletic Afrotheria. As the oldest record of an afrothere clade, identification of macroscelidean relatives in the North American Palaeocene argues against an African origin for Afrotheria, weakening support for linking placental diversification to the break-up of Gondwana.

Extraordinary fossorial adaptations in the oligocene palaeanodonts Epoicotherium and Xenocranium (Mammalia).

New fossils of the rare Oligocene mammals Xenocranium and Epoicotherium add information on their skulls and provide the first information on their postcranial skeletons. These epoicotheres, the latest surviving palaeanodonts, have numerous fossorial adaptations and must have been predominantly subterranean. Their skeletal specializations are similar to, and equal or surpass in degree of development, those of most living fossorial mammals. Principal modifications of the skull are the expanded, domed occiput with broad lambdoid crests, hypertrophy of the malleus-incus and related changes in other ear components, reduced eyes, and (in Xenocranium) a flaring, upturned, spatulate snout. The neck was strengthened by synostosis of the 2nd through 5th cervical vertebrae. The forelimb elements have exaggerated crests, processes, and fossae for muscles used in digging or in stabilizing certain joints. The scapula has a high, stout spine with bifid acromion, a "secondary spine," and an expanded postscapular fossa for attachment of the teres major muscle. The humerus has an elongate pectoral crest, large lesser tuberosity, long entepicondyle, and large hooklike supinator crest. The enormous incurved olecranon process of the ulna provided insertion for the massive triceps and origin for the carpal and digital flexors, and the latter gained mechanical advantage by incorporating in its tendon a large carpal sesamoid. In the greatly shortened hand, digit three is largest, with its metacarpal and proximal phalanx fused and its claw-bearing ungual-phalanx very large. These traits indicate that Xenocranium and Epoicotherium were among the most specialized "rapid-scratch" diggers ever to evolve. Their remarkable convergence to chrysochlorids reflects a similar mode of digging, with extensive use of the snout for loosening and lifting soil when making shallow foraging burrows. For deeper burrowing, the forelimbs probably loosened the soil while the rear limbs moved it behind. Like many extant subterranean mammals, Xenocranium and Epoicotherium were essentially sightless, but they were specialized for low frequency sound reception. Their extinction may have been due to a combination of environmental change and competition with other fossorial animals, such as proscalopine insectivores and rhineurid amphisbaenians.