Crystallite Orientation Discontinuities and the Evolution of Mammalian Enamel – Or, When is a Prism?

The nature and relationship of crystallite domains have been explored in fossil and extant enamels spanning an evolutionary period of 200 x 106 years. Minor crystallite orientation discontinuities, either linear or planar, were found to be consistent characteristics of all specimens examined. The earliest minor discontinuity is linear (convergence line), shown here in Oligokyphus and Eozostrodon. The convergence line would be the result of the occasional development of a conical Tomes\u27 process to the parent ameloblast. An increase in number and regularity of convergence lines, shown here in Haldanodon, marks the appearance of a regular pseudoprismatic enamel structure. The second minor discontinuity to appear is planar (seam), shown here in a dryolestid eupantothere. The seam has previously been deduced to relate developmentally to a central groove on the sloping floor-wall of the Tomes\u27 process pit. Coincident with the appearance of the seam is that of a rudimentary major planar discontinuity which does not enclose a domain to constitute what would normally be acknowledged as a prism. Its developmental basis would be the establishment of a steep wall and floor (however partial in circumference) to the Tomes\u27 process pit. The extent of the major planar discontinuities was found to increase subsequently to enclose a classically recognizable prismatic domain, shown here in Amphiperatherium, Hassianycteris, Smilodon and Felis. This would be consistent with the further development of a definitive floor and wall to the Tomes\u27 process pit. The sequential appearance of minor linear, minor planar and major planar discontinuities in crystallite orientation is seen as fundamental to the evolution of mammalian enamel structure

Cosmopolitanism among Gondwanan Late Cretaceous mammals

Consistent with geophysical evidence for the breaking up of Pangaea, it has been hypothesized that Cretaceous vertebrates on progressively isolated landmasses exhibit generally increasing levels of provincialism, with distinctly heightened endemism occurring at the beginning of the Late Cretaceous. The Cretaceous fossil record from the southern supercontinent of Gondwana has been much too poor to test this hypothesis with regards to mammals (Fig. 1). Early Cretaceous mammals are known only from isolated sites in Argentina, Australia, Cameroon and Morocco. Apart from several occurrences in South America, knowledge of Late Cretaceous Gondwanan mammals is limited to a single site in India that previously yielded a few specimens of placental mammals, and a site in Madagascar that previously yielded only one indeterminate tooth fragment. Here we report the occurrence of a highly specialized and distinctive group of extinct mammals, the Sudamericidae (Gondwanatheria), in the Late Cretaceous of Madagascar and India. These new records comprise the first evidence of gondwanatheres outside South America and the first indication of cosmopolitanism among Late Cretaceous Gondwanan mammals. Antarctica may have served as an important Cretaceous biogeographic link between South America and Indo-Madagascar

Digital reconstruction of the inner ear of Leptictidium auderiense (Leptictida, Mammalia) and North American leptictids reveals new insight into leptictidan locomotor agility

Leptictida are basal Paleocene to Oligocene eutherians from Europe and North America comprising species with highly specialized postcranial features including elongated hind limbs. Among them, the European Leptictidium was probably a bipedal runner or jumper. Because the semicircular canals of the inner ear are involved in detecting angular acceleration of the head, their morphometry can be used as a proxy to elucidate the agility in fossil mammals. Here we provide the first insight into inner ear anatomy and morphometry of Leptictida based on high-resolution computed tomography of a new specimen of Leptictidium auderiense from the middle Eocene Messel Pit (Germany) and specimens of the North American Leptictis and Palaeictops. The general morphology of the bony labyrinth reveals several plesiomorphic mammalian features, such as a secondary crus commune. Leptictidium is derived from the leptictidan groundplan in lacking the secondary bony lamina and having proportionally larger semicircular canals than the leptictids under study. Our estimations reveal that Leptictidium was a very agile animal with agility score values (4.6 and 5.5, respectively) comparable to Macroscelidea and extant bipedal saltatory placentals. Leptictis and Palaeictops have lower agility scores (3.4 to 4.1), which correspond to the more generalized types of locomotion (e.g., terrestrial, cursorial) of most extant mammals. In contrast, the angular velocity magnitude predicted from semicircular canal angles supports a conflicting pattern of agility among leptictidans, but the significance of these differences might be challenged when more is known about intraspecific variation and the pattern of semicircular canal angles in non-primate mammals

New Protocetid Whale from the Middle Eocene of Pakistan: Birth on Land, Precocial Development, and Sexual Dimorphism

BACKGROUND: Protocetidae are middle Eocene (49-37 Ma) archaeocete predators ancestral to later whales. They are found in marine sedimentary rocks, but retain four legs and were not yet fully aquatic. Protocetids have been interpreted as amphibious, feeding in the sea but returning to land to rest. METHODOLOGY/PRINCIPAL FINDINGS: Two adult skeletons of a new 2.6 meter long protocetid, Maiacetus inuus, are described from the early middle Eocene Habib Rahi Formation of Pakistan. M. inuus differs from contemporary archaic whales in having a fused mandibular symphysis, distinctive astragalus bones in the ankle, and a less hind-limb dominated postcranial skeleton. One adult skeleton is female and bears the skull and partial skeleton of a single large near-term fetus. The fetal skeleton is positioned for head-first delivery, which typifies land mammals but not extant whales, evidence that birth took place on land. The fetal skeleton has permanent first molars well mineralized, which indicates precocial development at birth. Precocial development, with attendant size and mobility, were as critical for survival of a neonate at the land-sea interface in the Eocene as they are today. The second adult skeleton is the most complete known for a protocetid. The vertebral column, preserved in articulation, has 7 cervicals, 13 thoracics, 6 lumbars, 4 sacrals, and 21 caudals. All four limbs are preserved with hands and feet. This adult is 12% larger in linear dimensions than the female skeleton, on average, has canine teeth that are 20% larger, and is interpreted as male. Moderate sexual dimorphism indicates limited male-male competition during breeding, which in turn suggests little aggregation of food or shelter in the environment inhabited by protocetids. CONCLUSIONS/SIGNIFICANCE: Discovery of a near-term fetus positioned for head-first delivery provides important evidence that early protocetid whales gave birth on land. This is consistent with skeletal morphology enabling Maiacetus to support its weight on land and corroborates previous ideas that protocetids were amphibious. Specimens this complete are virtual 'Rosetta stones' providing insight into functional capabilities and life history of extinct animals that cannot be gained any other way

Evidence for a Grooming Claw in a North American Adapiform Primate: Implications for Anthropoid Origins

Among fossil primates, the Eocene adapiforms have been suggested as the closest relatives of living anthropoids (monkeys, apes, and humans). Central to this argument is the form of the second pedal digit. Extant strepsirrhines and tarsiers possess a grooming claw on this digit, while most anthropoids have a nail. While controversial, the possible presence of a nail in certain European adapiforms has been considered evidence for anthropoid affinities. Skeletons preserved well enough to test this idea have been lacking for North American adapiforms. Here, we document and quantitatively analyze, for the first time, a dentally associated skeleton of Notharctus tenebrosus from the early Eocene of Wyoming that preserves the complete bones of digit II in semi-articulation. Utilizing twelve shape variables, we compare the distal phalanges of Notharctus tenebrosus to those of extant primates that bear nails (n = 21), tegulae (n = 4), and grooming claws (n = 10), and those of non-primates that bear claws (n = 7). Quantitative analyses demonstrate that Notharctus tenebrosus possessed a grooming claw with a surprisingly well-developed apical tuft on its second pedal digit. The presence of a wide apical tuft on the pedal digit II of Notharctus tenebrosus may reflect intermediate morphology between a typical grooming claw and a nail, which is consistent with the recent hypothesis that loss of a grooming claw occurred in a clade containing adapiforms (e.g. Darwinius masillae) and anthropoids. However, a cladistic analysis including newly documented morphologies and thorough representation of characters acknowledged to have states constituting strepsirrhine, haplorhine, and anthropoid synapomorphies groups Notharctus tenebrosus and Darwinius masillae with extant strepsirrhines rather than haplorhines suggesting that the form of pedal digit II reflects substantial homoplasy during the course of early primate evolution