Levels of Complexity in the Microstructure of Mammalian Enamel and Their Application in Studies of Systematics

Description and analysis of the complex structure of enamel can be facilitated through the application of a system of hierarchical levels of structural complexity. Five interdependent levels are distinguished. These are the levels of: 1) crystallites, 2) prisms, 3) enamel types, 4) schmelzmuster, and 5) dentition. This system provides a basis for analysis of both variation of particular structures and variation of structural types throughout a mammal\u27s dentition. Optimally, in wide ranging systematic and biomechanical studies, all levels of structural complexity should be considered, but lack of information about one level does not prevent significant analyses at other levels

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

Characteristics of the transverse 2D uniserial arrangement of rows of decussating enamel rods in the inner enamel layer of mouse mandibular incisors

The 2D arrangement of rows of enamel rods with alternating (decussating) tilt angles across the thickness of the inner layer in rat and mouse incisor enamel is well known and assumed to occur in a uniform and repetitive pattern. Some irregularities in the arrangement of rows have been reported, but no detailed investigation of row structure across the entire inner enamel layer currently exists. This investigation was undertaken to determine if the global row pattern in mouse mandibular incisor enamel is predominately regular in nature with only occasional anomalies or if rows of enamel rods have more spatial complexity than previously suspected. The data from this investigation indicate that rows of enamel rods are highly variable in length and have complex transverse arrangements across the width and thickness of the inner enamel layer. The majority of rows are short or medium in length, with 87% having < 100 rods per row. The remaining 13% are long rows (with 100-233 rods per row) that contain 46% of all enamel rods seen in transverse sections. Variable numbers of rows were associated with the lateral, central and mesial regions of the enamel layer. Each region contained different ratios of short, medium and long rows. A variety of relationships was found along the transverse length of rows in each region, including uniform associations of alternating rod tilts between neighboring rows, and instances where two rows having the same rod tilt were paired for variable distances then moved apart to accommodate rows of opposite tilt. Sometimes a row appeared to branch into two rows with the same tilt, or conversely where two rows merged into one row depending upon the mesial-to-lateral direction in which the row was viewed. Some rows showed both pairing and branching/merging along their length. These tended to be among the longest rows identified, and they often crossed the central region with extensions into the lateral and mesial regions. The most frequent row arrangement was a row of petite length nestled at the side of another row having the same rod tilt (30% of all rows). These were termed -focal stacks- and may relate to the evolution of uniserial rat and mouse incisor enamel from a multilayered ancestor. The mesial and lateral endpoints of rows also showed complex arrangements with the dentinoenamel junction (DEJ), the inner enamel layer itself, and the boundary area to the outer enamel layer. It was concluded that the diversity in row lengths and various spatial arrangements both within and between rows across the transverse plane provides a method to interlock the enamel layer across each region and keep the enamel layer compact relative to the curving DEJ surface. The uniserial pattern for rows in mouse mandibular incisors is not uniform, but diverse and very complex.Enamel rods in rodent incisors are arranged as stacked rows with alternating tilt angles across the transverse plane of the tooth. This study quantifies row patterns to determine if the layering of the rows and the alternating tilt angles are uniform or more irregular and complex in arrangement. The results indicated considerable diversity in row lengths, row tilts and 2D spatial row arrangements both within and between rows.Peer Reviewedhttps://deepblue.lib.umich.edu/bitstream/2027.42/151964/1/joa13053_am.pdfhttps://deepblue.lib.umich.edu/bitstream/2027.42/151964/2/joa13053.pd

New mammutids (Proboscidea) from the Clarendonian and Hemphillian of Oregon - a survey of Mio-Pliocene mammutids from North America

72 pages.A survey of Miocene-Pliocene Mammutidae from North America is provided. Two important but undescribed specimens from Unity (Clarendonian) and Hermiston (Hemphillian) are reported, and a previously described mandible from Black Butte (Clarendonian) is revisited. The Unity specimen, a maxilla with a tusked longirostrine mandible, represents Zygolophodon proavus. Upper and lower tusks are well developed. Mammutid species are best characterized by traits of the mandible, the upper tusks and lower and upper dentitions. However, they are rarely preserved in a single individual, as in the Unity specimen. A second mammutid mandible from the nearby, stratigraphically similar Black Butte locality has a short, tuskless symphysis and was described as “Mammut (Pliomastodon) furlongi”. We question the independence of this species, because both types of mandibles are coeval in several Eurasian localities. The widespread geographic and stratigraphic co-occurrence of different mandible types cannot be explained through a phylogenetic model alone. Therefore, we discuss whether these parallel mammutid occurrences can be interpreted as resulting from species diversity, intraspecific variability, pathological abnormities, or as a sex-linked morphological difference. The Hermiston maxilla from the upper Hemphillian represents Mammut matthewi. The large upper tusk differs from Z. proavus. The morphology of the mandible, however, remains unknown, thus its precise evolutionary stage is uncertain. The maxillary molars of the Hermiston specimen are larger than in Z. proavus however, we observe a high degree of variability in the size of mammutid molars so this character alone is not indicative of a specific evolutionary stage. The relatively small number of localities with informative specimens and the conservative dental morphology of mammutids throughout the Miocene and Pliocene obscures the potential occurrence of synchronous North American taxa. Our results favor the interpretation of a continuous and endemic evolution from Zygolophodon to Mammut and make an independent immigration of the genus Mammut from Eurasia less probable. Therefore, the genus name Mammut should be avoided for Eurasian mammutids

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

The Development of Cephalic Armor in The Tokay Gecko (Squamata: Gekkonidae: \u3cem\u3eGekko gecko\u3c/em\u3e)

Armored skin resulting from the presence of bony dermal structures, osteoderms, is an exceptional phenotype in gekkotans (geckos and flap-footed lizards) only known to occur in three genera: Geckolepis, Gekko, and Tarentola. The Tokay gecko (Gekko gecko LINNAEUS 1758) is among the best-studied geckos due to its large size and wide range of occurrence, and although cranial dermal bone development has previously been investigated, details of osteoderm development along a size gradient remain less well-known. Likewise, a comparative survey of additional species within the broader Gekko clade to determine the uniqueness of this trait has not yet been completed. Here, we studied a large sample of gekkotans (38 spp.), including 18 specimens of G. gecko, using X-rays and high-resolution computed tomography for visualizing and quantifying the dermal armor in situ. Results from this survey confirm the presence of osteoderms in a second species within this genus, Gekko reevesii GRAY 1831, which exhibits discordance in timing and pattern of osteoderm development when compared with its sister taxon, G. gecko. We discuss the developmental sequence of osteoderms in these two species and explore in detail the formation and functionality of these enigmatic dermal ossifications. Finally, we conducted a comparative analysis of endolymphatic sacs in a wide array of gekkotans to explore previous ideas regarding the role of osteoderms as calcium reservoirs. We found that G. gecko and other gecko species with osteoderms have highly enlarged endolymphatic sacs relative to their body size, when compared to species without osteoderms, which implies that these membranous structures might fulfill a major role of calcium storage even in species with osteoderms

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

A distinguishing feature of Pongo upper molars and its implications for the taxonomic identification of isolated hominid teeth from the Pleistocene of Asia

The taxonomic status of isolated hominoid teeth from the Asian Pleistocene has long been controversial due to difficulties distinguishing between pongine and hominin molars given their high degree of morphometrical variation and overlap. Here, we combine nonmetric and geometric morphometric data to document a dental pattern that appears to be taxonomically diagnostic among Pongo. We focus on the protoconule, a cuspule of well‐documented evolutionary history, as well as on shape differences of the mesial fovea of the upper molars