<Emphasis Type="Italic">Protungulatum</Emphasis>, Confirmed Cretaceous Occurrence of an Otherwise Paleocene Eutherian (Placental?) Mammal

Neither pre-Cenozoic crown eutherian mammals (placentals) nor archaic ungulates (“condylarths”) are known with certainty based on the fossil record. Herein we report a new species of the Paleocene archaic ungulate (“condylarth”) Protungulatum from undisputed Late Cretaceous aged rocks in Montana USA based on an isolated last upper premolar, indicating rare representatives of these common early Tertiary mammals appeared in North America a minimum of 300 k  years before the extinction of non-avian dinosaurs. The other 1200 mammal specimens from the locality are characteristic Late Cretaceous taxa. This discovery overturns the current hypothesis that archaic ungulates did not appear in North America until after the Cretaceous/Tertiary (K/T) boundary and also suggests that other reports of North American Late Cretaceous archaic ungulates may be correct. Recent studies, including ours, cannot determine whether Protungulatum does or does not belong to the crown clade Placentalia.     

A Systematic Study on Tooth Enamel Microstructures of Lambdopsalis bulla (Multituberculate,Mammalia) - Implications for Multituberculate Biology and Phylogeny

Tooth enamel microstructure is a reliable and widely used indicator of dietary interpretations and data for phylogenetic reconstruction, if all levels of variability are investigated. It is usually difficult to have a thorough examination at all levels of enamel structures for any mammals, especially for the early mammals, which are commonly represented by sparse specimens. Because of the random preservation of specimens, enamel microstructures from different teeth in various species are often compared. There are few examples that convincingly show intraspecific variation of tooth enamel microstructure in full dentition of a species, including multituberculates. Here we present a systematic survey of tooth enamel microstructures of Lambdopsalis bulla, a taeniolabidoid multituberculate from the Late Paleocene Nomogen Formation, Inner Mongolia. We examined enamel structures at all hierarchical levels. The samples are treated differently in section orientations and acid preparation and examined using different imaging methods. The results show that, except for preparation artifacts, the crystallites, enamel types, Schmelzmuster and dentition types of Lambdopsalis are relatively consistent in all permanent teeth, but the prism type, including the prism shape, size and density, may vary in different portions of a single tooth or among different teeth of an individual animal. The most common Schmelzmuster of the permanent teeth in Lambdopsalis is a combination of radial enamel in the inner and middle layers, aprismatic enamel in the outer layer, and irregular decussations in tooth crown area with great curvature. The prism seam is another comparably stable characteristic that may be a useful feature for multituberculate taxonomy. The systematic documentation of enamel structures in Lambdopsalis may be generalized for the enamel microstructure study, and thus for taxonomy and phylogenetic reconstruction, of multituberculates and even informative for the enamel study of other early mammals.     

The evolution of dinosaur tooth enamel microstructure

The evolution of tooth enamel microstructure in both extinct and extant mammalian groups has been extensively documented, but is poorly known in reptiles, including dinosaurs. Previous intensive sampling of dinosaur tooth enamel microstructure revealed that: (1) the three‐dimensional arrangement of enamel types and features within a tooth—the schmelzmuster—is most useful in diagnosing dinosaur clades at or around the family level; (2) enamel microstructure complexity is correlated with tooth morphology complexity and not necessarily with phylogenetic position; and (3) there is a large amount of homoplasy within Theropoda but much less within Ornithischia. In this study, the examination of the enamel microstructure of 28 additional dinosaur taxa fills in taxonomic gaps of previous studies and reinforces the aforementioned conclusions. Additionally, these new specimens reveal that within clades such as Sauropodomorpha, Neotheropoda, and Euornithopoda, the more basal taxa have simpler enamel that is a precursor to the more complex enamel of more derived taxa and that schmelzmusters evolve in a stepwise fashion. In the particularly well‐sampled clade of Euornithopoda, correlations between the evolution of dental and enamel characters could be drawn. The ancestral schmelzmuster for Genasauria remains ambiguous due to the dearth of basal ornithischian teeth available for study. These new specimens provide new insights into the evolution of tooth enamel microstructure in dinosaurs, emphasizing the importance of thorough sampling within broadly inclusive clades, especially among their more basal members.     

The oldest mammals from Antarctica,early Eocene of the La Meseta Formation,Seymour Island

New fossil mammals found at the base of Acantilados II Allomember of the La Meseta Formation, from the early Eocene (Ypresian) of Seymour Island, represent the oldest evidence of this group in Antarctica. Two specimens are here described; the first belongs to a talonid portion of a lower right molar assigned to the sparnotheriodontid litoptern Notiolofos sp. cf. N. arquinotiensis. Sparnotheriodontid were medium‐ to large‐sized ungulates, with a wide distribution in the Eocene of South America and Antarctica. The second specimen is an intermediate phalanx referred to an indeterminate Eutheria, probably a South American native ungulate. These Antarctic findings in sediments of 55.3 Ma query the minimum age needed for terrestrial mammals to spread from South America to Antarctica, which should have occurred before the final break‐up of Gondwana. This event involves the disappearance of the land bridge formed by the Weddellian Isthmus, which connected West Antarctica and southern South America from the Late Cretaceous until sometime in the earliest Palaeogene.     

ADDITIONAL SPECIMENS OF SUDAMERICID (GONDWANATHERIA) MAMMALS FROM THE EARLY PALEOCENE OF ARGENTINA

Abstract: The extinct, Cretaceous–Paleogene Gondwanatherians have previously been considered to be early xenarthrans, multituberculates and more recently Mammalia incertae sedis. However, the phylogenetic relationships of Gondwanatheria have yet to be resolved. In this paper, additional dental specimens of the gondwanatherian Sudamerica ameghinoi from the Early Paleocene Salamanca Formation of Argentina are described. These specimens provide additional information on Gondwanatheria affinities, sudamericid morphology and help support earlier hypotheses on Sudamerica dental formula and tooth categories. Sudamericid dental functional morphology and body mass estimates, based on measurements of isolated teeth, are inferred. Dental morphology such as hypsodonty, enamel microstructure and crown features do support a robust clade for Sudamericidae.     

First record of Entelodontidae (Mammalia,Artiodactyla) from the late Eocene of Southeast Asia

We describe two entelodontid upper premolars that were recovered from the late Eocene of the Krabi coal mine in southern Thailand. The size and morphology of the material suggest that it can be referred to Entelodon aff. E. gobiensis, a species known from the late Eocene to the early Oligocene of northern Asia and southern China. The Thai material documents for the first time the southernmost occurrence of entelodontids in Asia during the Paleogene and also suggests that Eocene Southeast Asian mammal localities might potentially yield further entelodontid remains mostly associated with selenodont ungulates.     

A new large philisid (Mammalia,Chiroptera, Vespertilionoidea) from the late Early Eocene of Chambi,Tunisia

Abstract: Among the new dental remains from the late Early Eocene of Chambi (Kasserine area, Tunisia) is a large‐sized upper molar of a new bat species, Witwatia sigei nov. sp. (Chiroptera, Vespertilionoidea, Philisidae), described herein. The locality of Chambi has revealed evidence for an early appearance of two modern microchiropteran superfamilies in Africa: Dizzya exsultans, a Philisidae, which is considered to be an archaic Vespertilionoidea, and an indeterminate Rhinolophoidea. In addition to D. exsultans, the new species, W. sigei, is the second representative of the Philisidae in this locality. W. sigei extends back to the late Early Eocene the occurrence of the genus Witwatia, which was previously only reported from the early Late Eocene of the Fayum (BQ‐2, Egypt). By analogy with the largest extant microbats, the large size of Witwatia suggests a tendency to the opportunistic diet of this taxon, thereby contrasting with the strict insectivory characterizing primitive bats found in other continents in the same epoch.     

The absence of muscle segment homeobox 2 leads to the pyroptosis of ameloblasts by inducing squamous epithelial hyperplasia in the enamel organ

Muscle segment homeobox 2 (MSX2) has been confirmed to be involved in the regulation of early tooth development. However, the role of MSX2 has not been fully elucidated in enamel development. To research the functions of MSX2 in enamel formation, we used a Msx2^−/− (KO) mouse model with no full Msx2 gene. In the present study, the dental appearance and enamel microstructure were detected by scanning electron microscopy and micro-computed tomography. The results showed that the absence of Msx2 resulted in enamel defects, leading to severe tooth wear in KO mice. To further investigate the mechanism behind the phenotype, we performed detailed histological analyses of the enamel organ in KO mice. We discovered that ameloblasts without Msx2 could secrete a small amount of enamel matrix protein in the early stage. However, the enamel epithelium occurred squamous epithelial hyperplasia and partial keratinization in the enamel organ during subsequent developmental stages. Ameloblasts depolarized and underwent pyroptosis. Overall, during the development of enamel, MSX2 affects the formation of enamel by regulating the function of epithelial cells in the enamel organ.     

Unique differentiation of radial enamel in Arsinoitherium (Embrithopoda,Tethytheria)

The enamel microstructure in molars of Arsinoitherium is reinvestigated and a new modification of radial enamel (RE), ‘arsinoitheriid radial enamel (ARE)’, is defined. It is characterised by alternating stripes with different organisation of the interprismatic matrix but no prism decussation. Recognition of this new subtype leads to a reinterpretation of structure previously identified as modified radial enamel and of Hunter–Schreger bands in Arsinoitherium. The newly differentiated ARE of Arsinoitherium is more derived in relation to corresponding microstructures of Palaeoamasia and Crivadiatherium. A careful reinvestigation of RE in other Paenungulata will be required to provide additional data bearing on phylogenetic reconstruction. The enamel of Phenacolophus argues against inclusion of this genus in the Embrithopoda.     

Prairie grass phytolith hardness and the evolution of ungulate hypsodonty

Silica phytoliths in grasses are thought to serve as a defence mechanism against grazing ungulates by causing excessive tooth wear. It is posited that they contributed to the evolution of hypsodonty in these animals. However, some have questioned whether grass phytoliths can abrade enamel. Here Mohs hardness testing was conducted on Blue Grama grass (Bouteloua gracilis) to determine phytolith hardness. Microindentation was performed on horse and American bison molars to establish dental constituent hardness values. To infer the phytoliths' abrasion capacity, the hardness values were contrasted. Phytolith hardness ranged from 18.0 to 191.5 HV. This is considerably softer than the values obtained for ungulate enamel, which range from 332.6 to 363.4 HV, but harder than the other dental constituents. Although Blue Grama phytoliths are incapable of directly abrading enamel, when viewed in conjunction with other data on phytolith hardness, there is considerable variation across grass species and some phytoliths are actually harder than ungulate enamel. Blue Grama grass phytoliths may even promote enamel wear due to pressure accentuation caused by the recession of softer tissues. Given these findings and considerations, it is plausible phytoliths served an integral role in the co-evolution of grasses and herbivorous ungulates, although more testing is needed to bear this out.     

Enamel Structure of Cuvieronius hyodon (Proboscidea, Gomphotheriidae) with a Discussion on Enamel Evolution in Elephantoids

Dental material of the South American elephantoid Cuvieronius hyodon from the Late Pleistocene of the Tarija Basin, Bolivia was sampled for a comprehensive analysis of the microstructure of the enamel. To examine variability at the dentition level, enamel samples of the upper incisor, second deciduous premolar, and molars were sectioned. The incisor and cheek teeth enamel is compared to that of other proboscideans in order to reveal phylogenetically and functional informative features useful to reconstruct the evolution of elephantoid enamel. Studies of the adaptations and evolution of proboscidean enamel have focused so far on molars. Nevertheless, given the possibility of an independent evolution of the enamel at different tooth positions, the variation of the enamel throughout the dentition needs to be taken into consideration when using enamel microstructural characters to infer proboscidean diversity and phylogeny. The results obtained from this study demonstrate the generality, among elephantoids, of the basic microstructural features of Cuvieronius hyodon enamel, allowing the characterization of the Elephantoid Enamel (EE). The differentiation between incisor and molar enamel seen in elephantoids is shown to represent a primitive elephantiform trait, as it also occurs in Phiomia. The three-layered enamel of the cheek teeth appears as the sole synapomorphy of the Elephantoidea, though the character might be homoplastic within the Proboscidea. Characters of the prisms cross-section might be used, on the other hand, to define less inclusive clades within the Elephantoidea.     

Enamel microstructure in lemuridae (mammalia,primates): Assessment of variability

This study describes the molar enamel microstructure of seven lemurid primates: Hapalemur griseus, Varecia variegata, Lemur catta, Lemur macaco, Lemur fulvus rufus, Lemur fulvus fulvus, and Lemur fulvus albifrons. Contrary to earlier accounts, which reported little or no prism decussation in lemurid enamel, both Lemur and Varecia molars contain a prominent inner layer of decussating prisms (Hunter-Schreger bands), in addition to an outer radial prism layer, and a thin, nonprismatic enamel surface layer. In contrast, Hapalemur enamel consists entirely of radial and, near the surface, nonprismatic enamel. In addition, for all species, prism packing patterns differ according to depth from the tooth surface, and for all species but Varecia (which also has the thinnest enamel of any lemurid), average prism area increases from the enamel-dentine junction to the surface; this may be a developmental solution to the problem of accommodating a larger outer surface area with enamel deposited from a fixed number of cells. Finally, contradicting some previous reports, Pattern 1 prisms predominate only in the most superficial prismatic enamel. In the deeper enamel, prism cross-sections include both closed (Pattern 1) and arc-shaped (Pattern 2 or, most commonly, Pattern 3). This sequence of depth-related pattern change is repeated in all taxa. It should also be emphasized that all taxa can exhibit all three prism patterns in their mature enamel. The high degree of quantitative and qualitative variation in prism size, shape, and packing suggests that these features should be used cautiously in phylogenetic studies. Hapalemur is distinguished from the other lemurids by unique, medially constricted or rectangular prism cross-sections at an intermediate depth and the absence of prism decussation, but, without further assessment of character polarity, these differences do not clarify lemurid phylogenetic relations. Some characters of enamel microstructure may represent synapomorphies of Lemuridae, or of clades within Lemuridae, but homoplasy is likely to be common. Homoplasy of enamel characters may reflect functional constraints. © 1994 Wiley-Liss, Inc.     

New Specimens of Reigitherium bunodontum from the Late Cretaceous La Colonia Formation,Patagonia, Argentina and Meridiolestidan Diversity in South America

We describe the first maxillae and additional new specimens of Reigitherium bunodontum, a small meridiolestidan from the Late Cretaceous La Colonia Formation, Patagonia, Argentina. The new material supports a dental formula of I?, C1, P4, M3, resolves postcanine positional uncertainty and corrects previous interpretations. Our phylogeny recovers Reigitherium as a meridiolestidan allied to other bunodont Mesungulatoidea, as the sister group of the Paleocene Peligrotherium. Posterior premolars/molars of Reigitherium, and to a smaller degree Peligrotherium, are dominated by an incomplete transverse ridge running between the protoconid-metaconid in the lowers and the paracone-stylocone in the uppers, semi-symmetrical basins developing mesially and distally from these central ridges. The trigonid-derived single transverse crest results from a mesial shift of the robust metaconid, an enhancement of the basin crest stretching from the protoconid/metaconid, and a shallower trigonid basin. The mesungulatoid condition, with its complete absence of talonid, contrasts sharply with that of therians with lophs, or transverse ridges, which involved at least one talonid-derived loph resulting in two transverse crests per tooth. Mesungulatoid meridiolestidans achieved complex tooth-on-tooth occlusion with a predicted increase in herbivory/omnivory, departing from the traditional sharp-cusp insectivores plesiomorphic for meridiolestidans and Mesozoic mammals in general. Reigitherium’s dramatic remodeling of the primitive meridiolestidan molar morphology, extensive continuous occlusal surface, accessory cuspules, and highly textured crenulated enamel illustrates one of most distinctive adaptations to herbivory among Mesozoic mammals.

     

Adenosine A2B receptor antagonist suppresses differentiation to regulatory T cells without suppressing activation of T cells

Neural crest cells (NCCs) are a multipotent embryonic cell population that contributes to the formation of various craniofacial structures including teeth. It has been generally believed that dental enamel is an ectodermal derivative, whereas the dentin–pulp complex and the surrounding supporting tissues originate from NCC-derived mesenchyme. These traditional concepts stem mainly from several early studies of fishes and amphibians. Recently, Wnt1-Cre/R26R mice, a mouse model for NCC lineage analysis, revealed the contribution of NCCs to mammalian tooth development. However, the discrepancy of expression patterns between different NCC-specific transgenic mouse lines makes it compulsory to revisit the cell lineage in mammalian tooth development. Here, we reevaluated the NCC lineage during mouse tooth development by using P0-Cre/R26R mice, another NCC-specific transgenic mouse line. Inconsistent with the traditional concepts, we observed the potential contribution of NCCs to developing enamel organ and enamel formation. We also demonstrated that the P0-Cre transgene was specifically expressed in migrating NCC in the hindbrain region, where NCC contributes to tooth, validating their applicability for NCC lineage analysis. Our unanticipated finding may change the general understanding of tooth development and provide new insights into dental stem cell biology.     

Postcanine microstructure in Cricodon metabolus,a Middle Triassic gomphodont cynodont from south‐eastern Africa

Cricodon metabolus is a trirachodontid cynodont from the Anisian (Middle Triassic) of eastern and southern Africa. It has labiolingually expanded (gomphodont) postcanines but also a sectorial tooth in the last postcanine locus. In this paper, we examine the crown microstructure of isolated sectorial and gomphodont postcanines belonging to the holotype specimen of this taxon using scanning electron microscopy. The enamel of both teeth is prismless and composed of discontinuous columnar divergence units, supporting the consistent presence of synapsid columnar enamel in cynognathians. Abundant tubules and numerous irregularly spaced incremental lines are also visible in the enamel and dentine layers in each tooth. This study reveals that the enamel thickness varies along the tooth row in Cricodon as the enamel layer of the gomphodont postcanines is 11.5 times thicker than that of the sectorial crown. It is likely that this difference reflects occlusal stresses and fewer replacements in gomphodont postcanines relative to sectorial teeth. Approximately 100 incremental growth lines of von Ebner are present in the dentine layer, indicating that the deposition of the dentine by odontoblasts occurred for three months before the animal's death.     

Tooth development in a scincid lizard, Chalcides viridanus (Squamata), with particular attention to enamel formation

Comparative analysis of tooth development in the main vertebrate lineages is needed to determine the various evolutionary routes leading to current dentition in living vertebrates. We have used light, scanning and transmission electron microscopy to study tooth morphology and the main stages of tooth development in the scincid lizard, Chalcides viridanus, viz., from late embryos to 6-year-old specimens of a laboratory-bred colony, and from early initiation stages to complete differentiation and attachment, including resorption and enamel formation. In C. viridanus, all teeth of a jaw have a similar morphology but tooth shape, size and orientation change during ontogeny, with a constant number of tooth positions. Tooth morphology changes from a simple smooth cone in the late embryo to the typical adult aspect of two cusps and several ridges via successive tooth replacement at every position. First-generation teeth are initiated by interaction between the oral epithelium and subjacent mesenchyme. The dental lamina of these teeth directly branches from the basal layer of the oral epithelium. On replacement-tooth initiation, the dental lamina spreads from the enamel organ of the previous tooth. The epithelial cell population, at the dental lamina extremity and near the bone support surface, proliferates and differentiates into the enamel organ, the inner (IDE) and outer dental epithelium being separated by stellate reticulum. IDE differentiates into ameloblasts, which produce enamel matrix components. In the region facing differentiating IDE, mesenchymal cells differentiate into dental papilla and give rise to odontoblasts, which first deposit a layer of predentin matrix. The first elements of the enamel matrix are then synthesised by ameloblasts. Matrix mineralisation starts in the upper region of the tooth (dentin then enamel). Enamel maturation begins once the enamel matrix layer is complete. Concomitantly, dental matrices are deposited towards the base of the dentin cone. Maturation of the enamel matrix progresses from top to base; dentin mineralisation proceeds centripetally from the dentin–enamel junction towards the pulp cavity. Tooth attachment is pleurodont and tooth replacement occurs from the lingual side from which the dentin cone of the functional teeth is resorbed. Resorption starts from a deeper region in adults than in juveniles. Our results lead us to conclude that tooth morphogenesis and differentiation in this lizard are similar to those described for mammalian teeth. However, Tomes processes and enamel prisms are absent.     

Origin and evolution of the Pseudorhyncocyonidae,a European Paleogene family of insectivorous placental mammals

Two new species of pseudorhyncocyonid, Fordonia lawsoni sp. nov. and Leptictidium prouti sp. nov. from the UK earliest Eocene, described here, are older than any previously recorded member of the family. They are represented by teeth from numerous loci, which allow a better understanding of the sparsely known dentitions of currently known pseudorhyncocyonids. This facilitates the recognition of two further species of Leptictidium, L. listeri sp. nov. from the Middle Eocene of Germany and L. storchi sp. nov. from the Late Eocene of France. Study of occlusal relationships also helps to fill gaps in our knowledge of missing tooth loci. Cladistic analysis of pseudorhyncocyonids with their previously judged closest relatives, the Leptictidae, Pantolesta and Palaeanodonta, shows that two European species, Diaphyodectes prolatus and Palaeictops? levei, formerly thought to be leptictids, are instead primitive pseudorhyncocyonids, extending the range of the family further back in time to the Middle Paleocene. P? levei is placed in the new genus Phakodon gen. nov. The analysis also shows that the Pseudorhyncocyonidae are sister group to the other three groups combined and that family‐level differentiation in this probable clade took place as early as the earliest Paleocene.     

Jaw mechanism and dental function in the late cretaceous basal eusuchian Iharkutosuchus

Iharkutosuchus makadii is a basal eusuchian crocodylian with multicusped teeth discovered from the Upper Cretaceous of Hungary. Skull and dentition morphology indicates an active food processing for this crocodylian. First among crocodylians, a combination of different analyses, including cranial adductor muscle reconstruction, tooth wear pattern, and enamel microstructure studies, is applied here to support this hypothesis. Data provide unambiguous evidence for significant dental occlusion that was a result of a unique, transverse mandibular movement. Reconstruction of the jaw adductors demonstrates strong muscles responsible for slow but active jaw closure as the motor of transverse jaw movement; nevertheless muscles producing rapid jaw closure were reduced. Macrowear orientations show a dominantly transverse movement of the mandibles completed by a slight anteroposterior component. Along with quadrate morphology, macrowear further indicates that this motion was accomplished by alternate rotation of the mandibles about the quadrate condyles. Dental morphology and wear patterns suggest two types of power stroke: a slicing–crushing stroke associated dominantly with anterior tooth–food–tooth contact (with a low degree of transverse mandibular movement) during in the early stage of mastication, and a grinding stroke with significant posterior tooth–tooth contact and a dynamic transverse movement occurring later. The patterns of microwear show a diverse diet for Iharkutosuchus including both soft and hard items. This is also supported by the microstructure of the thick, wrinkled enamel built up mostly by poorly developed columnar units. Based on wear patterns, ontogenetic variation in feeding habits of Iharkutosuchus is also recognized. J. Morphol., 2009. © 2009 Wiley‐Liss, Inc.