A new gomphodont cynodont (Traversodontidae) from the Middle–Late Triassic Dinodontosaurus Assemblage Zone of the Santa Maria Supersequence,Brazil

The tetrapod faunas from the terrestrial Middle–Late Triassic basins of Africa and South America are among the richest in the world, especially in non‐mammalian cynodonts. Despite the great abundance of cynodont specimens found in these basins, there are few known taxa that exhibit interbasinal distributions. Here we describe a new species of traversodontid cynodont of the genus Scalenodon from the Triassic Dinodontosaurus Assemblage Zone of the Santa Maria Supersequence, from the state of Rio Grande do Sul, Brazil. Scalenodon ribeiroae sp. nov. is based on a partial skull that possesses a combination of features not observed in any other South American traversodontid: ellipsoid upper postcanines with the transverse crest formed by three cusps, lacking a mesiobuccal accessory cusp, and with lingual cusp projected lingually creating a concave lingual surface on the upper postcanines; the paracanine fossa is positioned medially to the upper canine, and jugal lacks a suborbital process. A phylogenetic analysis places the new taxon in a basal position within the Family Traversodontidae, with the African Scalenodon angustifrons as sister‐taxon. The new specimen of Scalenodon represents the first record of this genus outside of the Manda Beds of Tanzania, and reinforces the biostratigraphical and biogeographical connection between Gondwanan Middle–Late Triassic tetrapod faunas. Although recent advances have been made, our current knowledge of these faunas is limited by the lack of absolute dates for most units and by uncertainties in the taxonomy and stratigraphical provenance of key fossils.     

New traversodontid (Therapsida: Cynodontia) teeth from the Upper Triassic of Habay-la-Vieille (southern Belgium)

Dwarf traversodontids (Therapsida, Cynodontia) are rare components of Late Triassic terrestrial faunas from western Europe.Habayia halbardieri n. gen., n. sp., described and discussed in the present paper, is based on one isolated upper postcanine from the Rhaetian of Habay-la-Vieille (southern Belgium). Another tiny tooth from the same locality is tentatively identified as a traversodontid lower postcanine. Dwarf cynodonts were well diversified in western Europe at the end of the Triassic. Their radiation, coupled with the sudden origin of mammals, corresponds to the development of the Rhaetian transgression which transformed western Europe into an archipelago and led to a reorganization of the terrestrial faunas.     

Geometric morphometric 3D shape analysis and covariation of human mandibular and maxillary first molars

Dental casts of 160 Greek subjects (80 males, 80 females) were scanned by a structured‐light scanner. The upper and lower right first molar occlusal surface 3D meshes were processed using geometric morphometric methods. A total of 265 and 274 curve and surface sliding semilandmarks were placed on the upper and lower molar surfaces, respectively. Principal component analysis and partial least square analysis were performed to assess shape parameters. Molars tended to vary between an elongated and a more square form. The first two principal components (PCs), comprising almost 1/3 of molar shape variation, were related to mesiodistal–buccolingual ratios and relative cusp position. Distal cusps displayed the greatest shape variability. Molars of males were larger than those of females (2.8 and 3.2% for upper and lower molars respectively), but no shape dimorphism was observed. Upper and lower molar sizes were significantly correlated (r² = 0.689). Allometry was observed for both teeth. Larger lower molars were associated with shorter cusps, expansion of the distal cusp, and constriction of the mesial cusps (predicted variance 3.25%). Upper molars displayed weaker allometry (predicted variance 1.59%). Upper and lower molar shape covariation proved significant (RV = 17.26%, P < 0.0001). The main parameter of molar covariation in partial least square axis 1, contributing to 30% of total covariation, was cusp height, in contrast to the primary variability traits exhibited by PC1 and PC2. The aim of this study was to evaluate shape variation and covariation, including allometry and sexual dimorphism, of maxillary and mandibular first permanent molar occlusal surfaces. Am J Phys Anthropol 152:186–196, 2013. © 2013 Wiley Periodicals, Inc.     

Evolution of the Tribosphenic Molar Pattern in Early Mammals,with Comments on the “Dual-Origin” Hypothesis

Development of the tribosphenic molar was a fundamental event that likely influenced the rise of modern mammals. This multi-functional complex combined shearing and grinding in a single chewing stroke, and provided the base morphology for the later evolution of the myriad dental morphologies employed by mammals today. Here a series of morphotypes are presented that represent stepwise acquisition of characters of the molar crown, in an effort to clarify homologies and functional analogies among molars of tribosphenic and tribosphenic-like mammals, as well as their putative sister groups. This is accomplished by evaluation of wear features, which provide direct evidence of occlusal function, and mapping these features on molars of the various morphotypes demonstrates their utility in determining homology. The original singular lower molar talonid cusp is homologous with the hypoconid, and upper molar cusp C in early mammals is homologous with the metacone (cusp “C” is a neomorph with variable occurrence). The lingual translation of the metacone to a position more directly distal to the paracone (as in Peramus) creates an embrasure for the lower molar hypoconid, and is accompanied by the development of the hypoconulid and a new shearing surface. Lastly, the Gondwanan radiation of tribosphenic-like mammals, the Australosphenida (including monotremes), is determined to be functionally non-tribosphenic. The Tribosphenida are restricted to Laurasian taxa, with an origin at or just prior to the Jurassic-Cretaceous boundary.     

Occlusal molar surfaces in females with Turner's syndrome

The aim of this study was to identify the molar occlusal features in 73 subjects with the Turner's syndrome (TS) and compared to a control group (CG) of 322 healthy females. The occlusal features were scored on dental plaster casts using the Scoring Procedures for Key Morphological Traits of the Permanent Dentition: The Arizona State University Dental Anthropology System (ASU). The results were analyzed through frequency, percentage and chi 2-test. TS subjects have more frequent reduction of cusp number, distolingual cusp on the upper molars and distal cusp on the lower molar, with the consequent reduction of the occlusal surface. Reduced size of occlusal surface and number cusps on upper molars resulted in the transformation of rhomboid occlusal shape into triangular, with the consequent loss of H-shaped groove system (in the upper right first molars H-shaped groove system was significantly less frequently found in TS (p < 0.05); in the upper left second molars H-shaped groove system was significantly less frequently found in TS (p < 0.01). The X-chromosome aneuploidy can cause a decrease in developmental homeostasis, which results in the alteration of apposition of the enamel and in consequently substantial changes of the molar occlusal morphological features.     

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.     

Dental Morphology of the Jurassic Holotherian Mammal Amphitherium, with a Discussion of the Evolution of Mammalian Post-Canine Dental Formulae

Four fragmentary mandibles from the Stonesfield Slate facies, Taynton Limestone Formation, Middle Bathonian (Middle Jurassic), England, represent two species of Amphitherium A. prevostii A. rixoni sp. nov. Both species had five lower premolariform teeth. The composite formula for the lower dentition of Amphitherium appears to have been I/4, C/1, P/5, M/6–7. The seventh molar is not present in the presumably oldest individual, and its presence is regarded as probably an individual variation.   The mosaic evolution of patterns of differentiation of the postcanine dentition from nonmammalian cynodonts to modern therian mammals is reviewed. It is concluded that Amphitherium probably had reached the grade of modern therians in the division between diphyodont premolars and monophyodont molars. The common ancestor of Amphitherium and zatheres probably had lost the primitive pattern of posterior shift of the postcanine dentition, which appears to have consisted of five premolars and four or possibly five molars. In Amphitherium the number of molars probably was secondarily increased.     

A traversodont cynodont from the Middle Triassic (Ladinian) of Baden-Württemberg (Germany)

Nanogomphodon wildi n. gen., n. sp. is based on a tiny lower postcanine tooth from the lower Lettenkeuper (Lower Keuper or Erfurt Formation; Ladinian) of Michelbach an der Bilz (Baden-Württemberg). It represents the first record of a traversodont cynodont from the Middle Triassic of Europe and exhibits a distinctive combination of dental features. Along with recent discoveries of other traversodont taxa from the Upper Triassic of eastern North America,Nanogomphodon indicates the existence of a distinct lineage of these cynodonts in the Northern Hemisphere.      

A new traversodontid cynodont from the Santa Maria Formation (Ladinian-Carnian) of southern Brazil, with a phylogenetic analysis of Gondwanan traversodontids

The Traversodontidae is a Triassic family of nonmammalian cynodonts, recorded worldwide, characterized by its bucco-lingually expanded postcanines. A recently discovered fauna in the locality of Santa Cruz do Sul, corresponding to the Santa Maria Formation of the Brazilian Middle/Upper Triassic (Ladinian/Carnian), is made up exclusively of nonmammalian cynodonts with an abundance of traversodontids. A new taxon, Santacruzodon hopsoni gen. et sp. nov . , the most commonly recorded traversodontid cynodont in the fauna, is described here. It is diagnosed by an autapomorphy, a very large posterolabial cusp in the upper postcanine, representing more than half of the labial crest, and a suite of traits present in other traversodontids. Among these features are the descendent flange of the jugal developed as a ball-shaped projection, also known in the Madagascan traversodontid Dadadon isaloi , and the presence of symmetrical incisors with numerous mesial and distal marginal cuspules, feature shared with the North American taxon Arctotraversodon plemmyridon . A phylogenetic analysis of traversodontids from Gondwana, based on a data matrix of 28 characters (mostly dental) and 15 terminals (13 traversodontids, including the new species, plus Diademodon and Trirachodon ) was performed. The new traversodontid appears as the sister taxon of Dadadon . A monophyletic Carnian group composed of Exaeretodon , Menadon and Scalenodontoides , with the Carnian Gomphodontosuchus , and the Ladinian Massetognathus as successive outgroups is also supported. In line with a recent phylogenetic hypothesis, a nonmonophyletic relationship for Scalenodon 'species' from the Anisian Manda beds also results from our analysis. Based on the cladogram obtained, morphological trends of change in the dentition of traversodontids throughout the Triassic are suggested.  © 2003 The Linnean Society of London, Zoological Journal of the Linnean Society, 2003, 139 , 529−545.     

Common solutions to resolve different dietary challenges in the ruminant dentition: The functionality of bovid postcanine teeth as a masticatory unit

Plasticity of tooth shape in mammals is of great adaptive value for the efficient exploitation of specific feeding niches and is a crucial mechanism for ecological diversification. In this study, we aimed to infer chewing effectiveness from the functional shape of different postcanine teeth within bovids, the most diverse extant group of large herbivorous mammals. We consider the postcanine dentition as a masticatory unit and test for differences related to food biomechanical properties, dietary abrasiveness, and chewing dynamics. We compare functional properties of the postcanine tooth row among species with well‐known dietary strategies by integrating digitalization of high‐resolution occlusal surface 3D‐models of upper postcanine dentitions and quantification of the indentation index (D), a structural parameter representing enamel complexity. We test for differences in the occlusal shape among tooth positions in the postcanine dentition using robust, heteroscedastic tests in a one‐way analysis of variance. Our results show three distinct patterns of enamel complexity along the tooth row: (1) D is more homogeneously distributed among tooth positions; (2) D increases gradually in the mesiodistal axis along the tooth row; and (3) D increases abruptly only at the transition between premolars and molars. We interpreted these patterns as different adaptive configurations of the postcanine tooth row relating to diet. Grass‐ and fruit‐eating bovids show the same abrupt increase in enamel complexity at the transition from premolars to molars. Intermediate feeding and leaf‐browsing species show the same gradual, mesiodistal increase in complexity along the tooth row. The absolute physical dietary resistance (biomechanical properties plus abrasiveness) and its relation to mechanical constraints of the chewing stroke are the likely selective factors leading to convergence of enamel complexity patterns along the tooth row among taxa with different diets. J. Morphol. 275:328–341, 2014. © 2013 Wiley Periodicals, Inc.     

Three-dimensional measurement of the occlusal surfaces of lower first molars of Australian aboriginals

The occlusal surfaces of lower first molars of Australian Aboriginals were measured in three dimensions with the aid of Moiré contourography. Molar cuspal heights in this population were higher than in Japanese (Mongoloid) but lower than in Dutch (Caucasoid) populations. Intercuspal distances were considerably larger than those in the two other populations. Populational differences in occlusal features may influence both craniofacial structures and jaw movements in the three populations. Low correlations between the cuspal heights and the intercuspal distances in the other two populations were also found in this population, indicating that human molar cuspal height is independent of the transverse size of the crown. Mean values for the height of the three principal cusps in lower first molars were less than those in upper ones. However, the height of the hypoconid, which was the highest in the lower molar cusps, showed almost the same mean value as the height of the upper three principal cusps, indicating that the height of the main functional cusp, in both upper and lower first molars, was almost the same.     

Prozostrodon brasiliensis,a probainognathian cynodont from the Late Triassic of Brazil: second record and improvements on its dental anatomy

Probainognathian cynodonts are conspicuous elements of the Assemblage Zones of the Triassic Santa Maria Supersequence in southern Brazil. Within this group, the derived clade Prozostrodontia, in which the crown group Mammalia is included, is taxonomic diverse in the Hyperodapedon and Riograndia AZs. We describe here the second known specimen (CAPPA/UFSM 0123) of Prozostrodon brasiliensis, until now only represented by its holotype. CAPPA/UFSM 0123 includes a right dentary with dentition. As in the holotype of P. brasiliensis, it has four lower incisors, pc4 with conspicuous cusp a, and small cusps b, c, and d, pc5-pc6 of ‘triconodont’ type with cusps a > c > b > d, with continuous lingual cingulum bearing up to six small discrete cusps, length of the lower tooth row more than half the length of the dentary, and relatively deep horizontal ramus of the dentary. The new specimen is about 25% smaller than the holotype and there is not a diastema between the canine and postcanine teeth, indicating its juvenile condition. Based on both known specimens of P. brasiliensis, a discussion on tooth replacement is presented, showing that the adult individual has more postcanine tooth morphotypes than the juvenile one.     

Diet and dental topography in pitheciine seed predators

Pitheciines (Pithecia, Chiropotes, and Cacajao) are a specialized clade of Neotropical seed predators that exhibit postcanine teeth with low and rounded cusps and highly crenulated occlusal surface enamel. Data on feeding ecology show that Pithecia consumes proportionally more leaves than other pitheciine species, and comparative studies demonstrate its greater molar relief and relative shearing potential. However, data on pitheciine food mechanics show that Pithecia masticates seeds with greater crushing resistance than those preferred by Chiropotes. This variation predicts an opposing morphology characterized by low and more rounded occlusal surfaces in Pithecia. We build on previous research using new methods for molar surface shape quantification by examining pitheciine second molar shearing crest length, occlusal relief, surface complexity, and surface curvature relative to nonseed specializing platyrrhines and within the context of the observed interspecific variation in pitheciine feeding ecology. Consistent with the previous analyses, our findings demonstrate that pitheciine molars exhibit low shearing, relief, and curvature compared with nonseed predators, independent of phylogeny. Pitheciines also exhibit highly “complex” occlusal topography that promotes the efficient breakdown of tough seed tissues. Overall, Pithecia, Chiropotes, and Cacajao share a similar topographic pattern, suggesting adaptation to foods with similar structural and/or mechanical properties. However, Cacajao differs in surface complexity, which reflects some variation in its feeding ecology. Contrary to the predictions, Pithecia and Chiropotes do not differ in any of the topographic variables examined. The range of demands imposed on the postcanine teeth of Pithecia might therefore select for an average topography, one that converges on that of Chiropotes. Am J Phys Anthropol, 2013. © 2012 Wiley Periodicals, Inc.     

The evolution of molar occlusion in the Cercopithecidae and early catarrhines

Those Eocene prosimians which are possible catarrhine ancestors have four blade-like crests on each lower molar. Each crest shears in sequence across two upper molar crests. Occluding crests are concavely curved to hold the foods being sheared. Each of two medial lower molar crests bordering the principal crushing surface shear past single upper molar crests at about the same time the lateral lower molar crests contact the second rank of upper molar crests. Grinding and crushing areas are restricted to hypoconid, trigonid, and protocone surfaces. Oligocene catarrhine molars have increased crushing-grinding capacities and maintained but modify their shearing. As the crushing surface of the protocone expands and a crushing hypocone is added, the “second rank” upper molar shearing crests are functionally reduced. At the same time medial crests are increasingly emphasized so that the total shearing capacity remains virtually unchanged. Marginal shearing blades are straight edged; leading edges of occluding blades are set at different angles to the occlusal plane so that blades contact at only one point at any given time. Early Primates have separate crushing basins surrounded by shearing blades. Catarrhines tend to link expanding crushing surfaces anteroposteriorly into a continuous surface between all molars. A cladistic analysis based on both new and previously recognized characters indicates that: 1, Apidium may be more closely related to Aegyptopithecus than to Parapithecus; 2, cercopithecids are derived from a Parapithecus-related stock; 3, Oreopithecus could equally well have come from an Apidium or Aegyptopithecus stock.     

New information about <Emphasis Type="Italic">Irajatherium hernandezi</Emphasis> Martinelli,Bonaparte, Schultz &; Rubert 2005 (Eucynodontia,Tritheledontidae) from the upper triassic (Caturrita Formation,Paraná Basin) of Brazil

In this contribution, new specimens of the tritheledontid eucynodont Irajatherium hernandezi, from the Late Triassic (Caturrita Formation) of southern Brazil, are analyzed. The new material provides significant information about incisor count, canine size and shape, basicranial morphology, and other previously unknown aspects of skull and dentition. A cladistic analysis with inclusion of the new data supports the assignment of Irajatherium to Tritheledontidae, basal to Chalimininae and Pachygenelinae. Previously unknown characters of Irajatherium revealed by the new material include: the presence of three lower incisors; the first lower incisor is enlarged; the presence of large upper and lower canines with deep paracanine fossa on the maxilla; almost complete upper and lower postcanine tooth row with a pattern similar to that of other tritheledontids (e.g. Pachygenelus and Chaliminia); there is a conspicuous crest on the inner surface of the maxilla for the attachment of the inferred maxillary turbinates; partially confluent jugular foramen and fenestra rotunda within the jugular fossa, separated by a finger-like projection of the posterolateral wall of the opisthotic; and hypoglossal foramina located outside the jugular fossa. Irajatherium is a key taxon for understanding the early evolution of ictidosaurs, a group of cynodonts closely related to mammaliaforms, during the cynodont–mammal transition from the Late Triassic to Early Jurassic.     

Evidence of strong stabilizing effects on the evolution of boreoeutherian (Mammalia) dental proportions

The dentition is an extremely important organ in mammals with variation in timing and sequence of eruption, crown morphology, and tooth size enabling a range of behavioral, dietary, and functional adaptations across the class. Within this suite of variable mammalian dental phenotypes, relative sizes of teeth reflect variation in the underlying genetic and developmental mechanisms. Two ratios of postcanine tooth lengths capture the relative size of premolars to molars (premolar–molar module, PMM), and among the three molars (molar module component, MMC), and are known to be heritable, independent of body size, and to vary significantly across primates. Here, we explore how these dental traits vary across mammals more broadly, focusing on terrestrial taxa in the clade of Boreoeutheria (Euarchontoglires and Laurasiatheria). We measured the postcanine teeth of N = 1,523 boreoeutherian mammals spanning six orders, 14 families, 36 genera, and 49 species to test hypotheses about associations between dental proportions and phylogenetic relatedness, diet, and life history in mammals. Boreoeutherian postcanine dental proportions sampled in this study carry conserved phylogenetic signal and are not associated with variation in diet. The incorporation of paleontological data provides further evidence that dental proportions may be slower to change than is dietary specialization. These results have implications for our understanding of dental variation and dietary adaptation in mammals.     

Locking Yourself Out: Diversity Among Dentally Zalambdodont Therian Mammals

We review the evolution of dental zalambdodonty across therian mammals. Among zalambdodonts, there is little or no occlusion between the protocone and talonid basin and one of the central cusps of the upper molars (metacone or paracone) and the talonid basin of the lower molars are lost or reduced. Over two dozen genera of therian mammals show zalambdodont molars, including tenrecids, chrysochlorids, Solenodon, the marsupial mole Notoryctes, the extinct placentals Apternodus, Oligoryctes, Parapternodus, and Koniaryctes, and the extinct marsupials Necrolestes and Yalkaparidon. The chiropteran Harpiocephalus is nearly zalambdodont. Transformation series provided by paleontological or ontogenetic data, plus occlusal relationships, can be used to determine the homology of molar cusps in zalambdodont taxa. Zalambdodont dental specialization does not appear to have led to ecological specialization. With the important exception of golden moles and tenrecs, dentally zalambdodont taxa are less speciose than their nonzalambdodont sister taxa.     

A new galesaurid (Therapsida: Cynodontia) from the Lower Triassic of South Africa

A new galesaurid cynodont, Progalesaurus lootsbergensis gen. et sp. nov., is described on the basis of a well-preserved skull, lower jaw, right scapula and left atlantal neural arch. Autapomorphies of Progalesaurus include postcanine teeth bearing numerous mesial and distal accessory cusps that flank a recurved main cusp, a post-temporal fenestra bordered by the squamosal ventrally and a large external naris. Progalesaurus is similar to Galesaurus in possessing a poorly defined masseteric fossa on the dentary, a strongly recurved main cusp of the postcanine dentition, an incomplete secondary palate and a similar basisphenoid-parasphenoid morphology. A cladistic analysis of ten early cynodont genera resolves a monophyletic Galesauridae encompassing Cynosaurus , Progalesaurus and Galesaurus , although support for this clade is weak. Procynosuchus and Dvinia are placed at the base of Cynodontia whereas Thrinaxodon and Platycraniellus are positioned higher, but outside of Eucynodontia. The holotype and only known specimen of Progalesaurus was collected during systematic prospecting of Permo/Triassic boundary strata at New Lootsberg Pass, Graaff-Reinet District, South Africa. The discovery of Progalesaurus increases the number of valid Early Triassic cynodonts to four and sheds light on the tempo of early cynodont diversification after the end-Permian mass extinction.     

A new tritylodontid from the Kayenta formation of Arizona

Kayentatherium wellesigen. sp. now is a new tritylodont (Thcrapsida: Synapsida) from the Kayenta Formation of Arizona, U.S.A. Only the skull and parts of the lower jaw are known. The first upper incisor is apparently absent, so the dentition consists of one upper incisor followed, after a diastema, by a row of postcanines, which are replaced posteriorly. The cusp formula of these upper postcanines is 2, 2 (+ a ) 2, a being a tiny mesial cusp. The mean breadth/length ratio of these teeth is 1.59. Kayentatherium is related to the Tritylodon Bienotherium lineage, but is considerably more advanced than the latter, which is of Upper Triassic Lower Jurassic age. This indicates that the Kayenta formation must be Lower or Middle Jurassic age, with agrees with the conclusions of Welles.