A unique form of dental bilophodonty and a functional interpretation of peculiarities in the masticatory system of Arsinoitherium (mammalia,Embrithopoda)

The highly autapomorphic upper molar bilophodonty of the Oligocene mammal, Arsinoitherium (Embrithopoda) is an extreme form of dilambdodonty effected by lingual positioning of normally buccally situated cusps with reduction of lingual cusps. This effectively limits the molar dentition to a single phase shearing occlusal motion. Molar and premolar morphology is very different, premolars exhibiting high longitudinal ectolophs and typical two phase occlusal morphology. A double faceted mandibular condyle and angular discontinuity between lower molar and premolar dentitions is interpreted as a means of separating premolar from molar occlusion. A bifunctional masticatory system is proposed whereby efficient premolar occlusion is achieved only after a repositioning of the temporomandibular joint. Loss of phase II occlusion in the molars is compensated by maintenance of a crushing/grinding mode in the premolars. This coupled with the ability to maintain high occlusal pressures along the length of the mandible explains the unbroken dental arcade. Arsinoitheres therefore possess an extremely specialised masticatory apparatus and are interpreted as highly selective browsing herbivores.     

The assessment of jaw movement direction from dental microwear

In order to resolve several controversies about unusual jaw movements such as thegosis and orthal retraction, information about direction of jaw movement is essential. Ryan (1979a,b) proposed that asymmetry of microwear striations might provide such data. In vivo occurrences of asymmetric striations on chimpanzee molars are presented and analyzed. Results suggest that either thegosis contacts commonly occur on chimpanzee molars or that Ryan's model, as developed in vitro, does not accurately apply to naturally occurring molar microwear.     

The ontogeny of premolar dental wear in Cercocebus albigena (cercopithecidae)

The orientation of striated wear facets on primate teeth serves as a useful guide for reconstructing jaw movements during mastication. Most wear facets on the molars are formed during one of the two well-documented movements, Phase I or Phase II, of the power stroke. Another jaw movement direction, “orthal retraction” (OR) has been proposed to account for a third set of facets occasionally present on the pointed tips of premolars and molars. Evidence advanced here indicates that OR facets on pointed anterior premolars (P₃) of cercopithecoids are actually Phase I facets that have become reoriented as a result of a rotation of this tooth during its eruption. “Orthal retraction” probably does not exist as a discrete masticatory phase.     

Genetic structure of a tribal population, the Yanomama Indians. XIII. Dental microdifferentiation.

Data are presented on the frequency of the following eight dental traits in 635 Yanomama and 65 Makiritare Indians: upper central incisor rotation or winging, shoveling of maxillary incisors, maxillary molar hypocone reduction, Carabelli's trait, mandibular molar cusp number, mandibular molar cusp pattern rotation of second lower premolar, and pattern of second lower premolar cusps. Yanomama dentition is unusual in the high frequency of six cusps on the mandibular molars. There is marked dental microdifferentiation between villages; significant agreement was observed between a matrix of pairwise "dental distances" based on six morphological traits and corresponding matrices based on 11 genetic systems and on geographic location.     

山西垣曲先炭兽类一新种

本文主要报道1983年9月,在山西省垣曲县寨里发现的一件较完整的先炭兽类老年个体上颌骨.带有颇为完整的左、右齿列. I~1-P~2 各齿前、后均有较长的齿隙. P~1、P~2、P~3 前的齿隙颌骨上,保留有乳前臼齿的齿槽.这块标本在大小及形态特征上,与目前所知先炭兽属 Anthracokeryx 中任何种都有相当大的差异.它代表了该属中的一个新种.     

Cercopithecoid canine tooth honing mechanisms

The form of the unworn male Cercopithecoid maxillary canine tooth (C') is effectively adapted for stabbing and slashing. Its essential features are maintained by wear against the mandibular canine (C₁) and first premolar (P₃) teeth. The cusp tip of C₁ is sharpened by reciprocal wear against C'. The distribution of apposing wear facets indicates that functional attrition results from honing activity largely distinct from mastication. Functional attrition also occurs in reduced form in females and is produced within the masticatory excursive range. The significance of the “sectorial” form of P₃ is analyzed. Its elongated mesiobuccal surface serves the dual purpose of honing the distal cutting edge of C' and functioning as a cutting block against which vegetation is stabilized and shredded by the cervical third of the distal cutting edge of C'. Behavioral aspects of honing are correlated with field observations linking tooth grinding with aggression, tension release, and communication. Parallel human behavior is cited and the suggestion is made that human tooth grinding with its highly charged emotional overtones is largely relict behavior that once had high survival value in a canine tooth honing context.     

Studies of fossil hyaenas: affinities of Lycyaenops rhomboideae Kretzoi from Pestlorinc, Hungary

The holotype of the lower-middle Pliocene hyaenid Lycyaenops rhomboideae is redescribed and compared with contemporaneous hyaenids of the genera Lycyaena, Hyaenictis, Chasmaporthetes and Pliocmcuta. These comparisons show that the material represents a valid and distinct genus and species. The genus Lycyaenops is referred to the Chasmaporthetes lineage on the basis of its two-cusped m 1 talonid with reduced entoconid, reduced posterolingual cingulum cusp on p4 and premolar accessory cusps set in a straight line. It is distinguished from all other genera of that lineage by its smaller premolar accessory cusps, broad premolars and squared-off and very broad posterior premolar shelves. The species L. silberbergi, previously assigned to Chasmaporthetes, is also referred to Lycyaenops. It differs from L. rhomboideae in its greater development of the premolar accessory cusps and less developed posterior premolar shelves, but shares the broad, squared-off premolars. The interrelationships of Hyaenictis, Chasmaporthetes and Lycyaenops are at present best described by an unresolved trichotomy, with Lycyaena as its sister taxon.     

Mastication in springhares,Pedetes capensis: A cineradiographic study

Analysis of lateral and dorsoventral radiographic films shows that ingestion, transport, and mastication in Pedetes capensis (Rodentia) are cyclic and their movement patterns are essentially similar for the three food types offered. During the ingestion cycle, closing of the mouth is accompanied by a backward translation of the condyles, so that movement is predominantly orthal. During the opening stage, the extent of the anterior condylar translation is smaller. As a result the mandibular incisors move ventrally and posteriorly. During the ingestion cycles, food is transported to the back of the tongue, with the transverse rugae and the folds of the upper lip playing important roles. Springhares show a bilateral masticatory pattern; food is chewed on both sides simultaneously. During chewing, the condyles lie in their most forward position at maximum opening of the mouth. The mouth is closed by rotation of the lower jaw around the temporomandibular joint coupled with posterior condylar translation. At the beginning of the slow-closing stage, the upward rotation of the mandible slows and the jaw slowly shifts forward. During the grinding stage, the mandible is shifted forward with both toothrows in occlusion. During the opening stage, the jaw returns to its starting position. Comparison of kinematic and anatomical data on rodent mastication suggests that some dental characteristics form the most important factors regulating the masticatory pattern and consequently allow reasonably reliable prediction of rodent masticatory patterns.     

MOLAR OCCLUSION IN LATE TRIASSIC MAMMALS

1. A new genus and species of late Triassic mammal, Megazostrodon rudnerae, from Lesotho in southern Africa is described. The molars are similar to those of the British Eozostrodon parvus except that they are slightly larger and the upper molars have a large external cingulum supporting well-developed cusps. 2. Molar occlusion is discussed in two groups of late Triassic mammals: Eoxostrodon and the closely related Megazostrodon on one the hand and the unnamed primitive symmetrodonts on the other. It is shown that in Eoxostrodon the upper and lower molars did not have matching occlusal surfaces upon eruption but that wear produced matching occlusal surfaces. These surfaces are confined to the internal surface of the upper molars and the external surface of the lower molars and form a series of wide-angled triangles. The main cusp of an upper molar occluded between the main and posterior subsidiary cusp of the lower molar and the main cusp of the lower molar occluded between the main and anterior subsidiary cusp of the upper molar, 3. It is shown that the molars of Docodon and HaIdanodon were possibly derived from those of a primitive mammal such as Eozostrodon. The transition involved the development on the upper molars of an internal extension which, as it increased in size, established contact with the dorsal surfaces of two adjacent lower molars. The process involved is fundamentally different from that leading to tribosphenic molars. 4. In Megaxostrodon the main cusp of the upper molars occluded between the posterior and anterior subsidiary cusps of two adjacent lower molars, i.e. more posteriorly than in Eozostrodon. Primitive Rhaetic symmetrodonts were derived from mammals which had this type of occlusion and which were also closely related to Eoxostrodon and Megaxostrodon. The transition involved a rotation of the subsidiary cusps of the upper molars externally and those of the lower molars internally. This rotation increased the shearing surfaces between occluding upper and lower molars. Cusp rotation was carried further in the acute-angled symmetrodonts (Peralestes and Spalacotherium) and pantotheres. It appears that marked cusp rotation was coupled with the acquisition of transverse movements of the lower jaw during mastication. Transverse movement was apparently not possible in cynodonts, in Eoxostrodon (and related forms) and in Docodon. 5. The evolution of therian molars involves cusp rotation as originally proposed by the Cope—Osborn theory. Criticisms of the Cope—Osborn theory are re-evaluated in light of the new late Triassic material. 6. In Rhaetic symmetrodonts, molar wear produces matching occlusal facets, but the amount of attrition necessary to produce these facets was considerably less than in Eoxostrodon. In acute-angled symmetrodonts and in pantotheres, the molars erupt with more precise occlusal surfaces and attrition was not necessary to produce matching surfaces. 7. On the basis of the structure of the molar teeth it was concluded that Eozostrodon, Megazostrodon and Erythrotherium were closely related to the Rhaetic symmetrodonts. Slightly different occlusal relationships between upper and lower molars indicated that in these early mammals constant occlusal relations were being established. 8. Primitive cynodonts, such as Thrinaxodon, are characterized by alternate tooth replacement; there is a total lack of a constant occlusal relationship between upper and lower postcanine teeth. In Thrinaxodon individual postcanines were replaced several times. The crown structures of successive generations of postcanines were different so that a freshly erupted postcanine tooth had a crown structure quite distinct from the tooth which it replaced. It has been shown that the crown structure of one of the generations of postcanine teeth of Thrinaxodon is almost identical to that of Eozostrodon except that Thrinaxodon postcanines have a single root, On the basis of this similarity and the over-all structure of the primitive cynodont skull, it was concluded that Rhaetic mammals (excluding ictidosaurs and haramyids) could be derived from primitive cynodonts. 9. All the orders of Jurassic mammals (with the possible exception of multituber-culates) were probably derived from late Triassic mammals. The apparent close relationship of late Triassic mammals is evidence of a monophyletic origin of this class.     

Differences in the Striaton Pattern in individuals with Unilateral Buccal Pathologies

A sample of individuals with different unilateral pathologies affecting the masticatory apparatus has been studied. Replicas of the same teeth (first or second molar) on both sides have been obtained and observed by SEM. The number, length and orientation of buccal striations have been determined for each individual. Differences in the microwear pattern have been observed between pathological sides. Each individual displays a particular striation pattern, especially referred to the striation number variables. It is concluded that individuals exhibiting pathologies likely to affect mastication should be excluded from studies relating striation patterns to diet.     

Der Bau der Canini bei den Paulchoffatiidae (Multituberculata; Ober-Jura)

Canines are preserved among Multituberculata only in the upper jaw of the Paulchoffatiidae, the Pinheirodontidae and the North American genusGliodon Engelmann &; Callison, 1999. They resemble the anterior premolars (p^1–3) in the morphology of their crown, but they differ from them by the presence of only one root. In the present paper, 126 isolated canines of the Paulchoffatiidae from the Guimarota coal-pit in Portugal are treated. They show a wide morphological variation, from bicuspid to pentacuspid expression, and they can be grouped into 18 morphological units (Tab. 1). The bicuspid canines (11 specimens) show one buccal and one lingual cusp, arranged side by side, the latter normally being larger than the former. In the tricuspid canines (32 specimens) the cusps (one buccal and two lingual) are arranged in a triangle, with a great variation in the position of the cusps to each other. Tetracuspid canines (69 specimens) are the dominating group. Two buccal and two lingual cusps are present, which differ markedly in their largeness and their position to each other. The teeth differ also much in the shape of their crown. The pentacuspid canines (14 specimens) show two buccal and three lingual cusps. The variation in the arrangement of the cusps and in the shape of the crown is similar as in the tetracuspid specimens. One or two small cuspules can be present at the anterior border of the crown additionally to the main cusps in the canines. Tricuspid and tetracuspid canines are present also in the skulls of the Paulchoffatiidae, whereas bicuspid and pentacuspid canines are known only as isolated specimens. In the Pinheirodontidae the canines are tricuspid or tetracuspid, inGlirodon they are unicuspid. The Paulchoffatiidae and Pinheirodontidae — Paulchoffatiid line sensuKielan-Jaworowska &; Hurum, 2001 — are characterized by increasing premolarization of the upper canines. With that they differ markedly from all other multituberculates where the canini become reduced.     

A geometric morphometric analysis of hominin upper premolars. Shape variation and morphological integration

This paper continues the series of articles initiated in 2006 that analyse hominin dental crown morphology by means of geometric morphometric techniques. The detailed study of both upper premolar occlusal morphologies in a comprehensive sample of hominin fossils, including those coming from the Gran Dolina-TD6 and Sima de los Huesos sites from Atapuerca, Spain, complement previous works on lower first and second premolars and upper first molars. A morphological gradient consisting of the change from asymmetric to symmetric upper premolars and a marked reduction of the lingual cusp in recent Homo species has been observed in both premolars. Although percentages of correct classification based on upper premolar morphologies are not very high, significant morphological differences between Neanderthals (and European middle Pleistocene fossils) and modern humans have been identified, especially in upper second premolars. The study of morphological integration between premolar morphologies reveals significant correlations that are weaker between upper premolars than between lower ones and significant correlations between antagonists. These results have important implications for understanding the genetic and functional factors underlying dental phenotypic variation and covariation.     

Molar occlusion and jaw mechanics of the Eocene primate Adapis

Wear facets on molars of the Eocene primate Adapis magnus are described. Striations on these wear facets indicate three separate directions of mandibular movement during mastication. One direction corresponds to a first stage of mastication involving orthal retraction of the mandible. The remaining two directions correspond to buccal and lingual phases of a second stage of mastication involving a transverse movement of the mandible. The mechanics of jaw adduction are analysed for both the orthal retraction and transverse stages of mastication. During the orthal retraction stage the greatest component of bite force is provided by the temporalis muscles acting directly against the food with the mandible functioning as a link rather than as a lever. A geometrical argument suggests that during the transverse stage of mastication bite force is provided by the temporalis muscles of both sides, the ipsilateral medial and lateral pterygoid muscles, and the contralateral masseter muscle.     

Wear pattern and functional morphology of dryolestoid molars (Mammalia, Cladotheria)

Pretribosphenic dryolestoid molars are characterized by a reversed triangular alignment of the “primary trigon” (formed by the paracone, metacone and stylocone) and trigonid crucial for the embrasure shearing process. These molars are abraded along the protocristid and paracristid, and show a typical wear pattern with mesially and distally sloping dentine fields due to their thin enamel. The wear pattern of lipotyphlan and didelphid tribosphenic molars with considerably thicker enamel does not show this sloping. In dryolestoid molars two directions of striations occur. Steeper striations oriented linguo-buccally are present on facet 1 below the protocristid, and about 10° less inclined striations of the same direction have been observed near the talonid base. This reflects the railing function of the hypoflexid for the paracone of the corresponding upper molar. Facet 3 in the hypoflexid gets steeper with progressive wear, whereas facets 1 and 2 on the mesial and distal sides of the trigonid are flattened during wear. In the masticatory process the hypoflexid has mainly a shearing function with a crushing component because of its lesser inclination than the functional shearing surfaces below the trigonid crests. Striations on the exposed dentine field along the paracristid and in the guiding groove of facet 3 indicate that these two surfaces were formed by attrition (tooth to tooth contact). The exposed dentine fields at the cusp apices and along the protocristid are gauged and therefore must be a result of abrasion (tooth to food contact).     

Evolutionary and developmental dynamics of the dentition in Muroidea and Dipodoidea (Rodentia, Mammalia)

SUMMARY When it comes to mouse evo‐devo, the fourth premolar–first molar (P4–M1) dental complex becomes a source of longstanding controversies among paleontologists and biologists. Muroidea possess only molar teeth but with additional mesial cusps on their M1. Developmental studies tend to demonstrate that the formation of such mesial cusps could result from the integration of a P4 germ into M1 during odontogenesis. Conversely, most Dipodoidea conserve their fourth upper premolars and those that lost these teeth can also bear additional mesial cusps on their first upper molars. The aim of this study is to assess this developmental model in both Muroidea and Dipodoidea by documenting the morphological evolution of the P4–M1 complex across 50 Ma. Fourteen extinct and extant species, including abnormal and mutant specimens were investigated. We found that, even if their dental evolutionary pathways strongly differ, Dipodoidea and Muroidea retain common developmental characteristics because some of them can present similar dental morphological trends. It also appears that the acquisition of a mesial cusp on M1 is independent from the loss of P4 in both superfamilies. Actually, the progressive decrease of the inhibitory effect of P4, consequent to its regression, could allow the M1 to lengthen and mesial cusps to grow in Muroidea. Apart from these developmental explanations, patternings of the mesial part of first molars are also deeply constrained by morpho‐functional requirements. As there is no obvious evidence of such mechanisms in Dipodoidea given their more variable dental morphologies, further developmental investigations are needed.     

Cusp size, sexual dimorphism, and heritability of cusp size in twins.

Overall measures of mandibular molars reflect the combined size contributions of the component cusps and ridges. Until now, the size hierarchy of primary and permanent mandibular molar cusps remained unclear. This paper utilizes the relative plane surface areas (basal area dimensions) of the individual molar cusps, as assays of cusp size to demonstrate cusp size variations within populations, antimere cuspal variations, sexual dimorphism, and, the heritability of cusp size. Duplicate dental casts from 199 pairs of like-sexed twins provide the raw dats. Defined anatomic landmarks on the occlusal surfaces were reduced to X-Y rectangular coordinates prior to the computation of the basal areas dimensions. The results establish a cusp size hierarchy specific for molar type, i.e., five-cusped molars with a distal fovea and distal marginal ridge (5fd), five-cusped molars without a distal fovea and without a distal marginal ridge (5o), and four-cusped molars (4c). Sexual dimorphism in cusp size is apparent in 5fd molar cusped but not in 5o molar cusps. However, males have a significantly higher frequency of 5fd molars. Females have a higher frequency of smaller 5o and 4c molars which have fewer crown components. Moreover, female 5o molars have cusps as large as or larger than 5o male molor cusps. Right-side-left-side differences exist between antimere cusps based on relatively low correlations. The mirroring of molor types occurs infrequently. When observed, most intrapair differences for cusp size, using F-ratios, indicate a low component of hereditary variability.     

Anterior dental cutting in the Laetolil hominids and the evolution of the bicuspid P3

The younger Laetolil hominids provide evidence of a unique anterior cutting complex with a chisel-like action occurring between the lingual-distal C face and a transverse ridge on the P₃ perpendicular to it, extending between the subequal cusps. An earlier adaptation to more efficient grinding may have resulted in reduced canine projection and the development of the lingual P₃ cusp, raising the ridge between it and the buccal cusp. This development acted to retain the anterior cutting function by shifting it to the top of the premolar.     

The dental morphology of Pima Indians

Fourteen morphologic crown traits were observed in a sample of 1528 Pima Indians of south-central Arizona. Pima dentitions are characterized by high frequencies of shoveling, incisor winging, the hypocone, the lower canine distal accessory ridge, cusp 6, and the protostylid. They exhibit low frequencies of the metaconule and lower premolar multiple lingual cusps and moderate frequencies of the canine tubercle, Carabelli's trait, cusp 7, and lower second molars with four cusps and X groove patterns. When Pima crown trait frequencies were compared to those of 13 Southwest Indian samples, their closest affinities were to other Uto-Aztecan groups, the Papago and Hopi. The Pima are most divergent from Athapaskans and are also clearly removed from Yuman speaking groups and the Zuni. In general, the pattern of dental morphologic variation in the Southwest corresponds closely to linguistic divisions.     

Tooth function and replacement in early Mesozoic ornithischian dinosaurs: implications for aestivation

Thulborn (1978, Leihaia II ) suggests that ornithischian dinosaurs of the upper Stormberg Series (Late Triassic-Early Jurassic) of southern Africa underwent aestivation during an annual dry season. His argument, based on an interpretation of tooth function and replacement in heterodontosaurids, is: (1) unequivocal evidence of tooth replacement is not seen, and (2) piecemeal replacement of the dentition would be incompatible with maintenance of a fore-aft grinding function of the teeth; therefore, the entire dentition must have been rapidly replaced as a unit during periods of non-feeding, i.e. during aestivation. However, study of tooth wear patterns in Lanasaurus, Lycorhinus , and Heterodontosaurus show that jaw movements during mastication were orthal (open-and-close) and lateral to medial, not forwards and backwards. Differences in degree of tooth wear would not interfere with masticatory movements. Patterns of differential wear indicate that tooth replacement was not periodic but continuous, as in other reptiles. Zahnreihen , with a Z-spacing of about 3.0, are recognizable. Replacement ceased in mature individuals. The dentition shows adaptations for prolonging its effective life despite heavy wear. Differential tooth wear is incompatible with the idea of replacement of the entire dentition as a unit during an hypothesized period of aestivation. Thulborn's suggestion of aestivation in fabrosaurid ornithischians is also shown to be unlikely.     

Phylogeny and Systematics of Multituberculate Mammals

We present a synopsis of high-rank multituberculate systematicsand a manually generated cladogram illustrating multituberculate interrelationships. We divide the Multituberculata into the paraphyletic suborder 'Plagiaulacida', an apparently monophyletic suborder Cimolodonta, and one family incertae sedis. Within 'Plagiaulacida' we recognise three informal lines: paulchoffatiid (three families), plagiaulacid (three families) and allodontid (two families and the genus Glirodon). The Cimolodonta are divided into an informal Paracimexomys group; three superfamilies: Ptilodontoidea, Djadochtatherioidea (new), and Taeniolabidoidea (restricted to Taeniolabididae); and five families (superfamily incertae sedis): Eucosmodontidae, Microcosmodontidae, Cimolodontidae, Boffiidae, and Kogaionidae; and some genera incertae sedis. New characters used in our analysis are (1) a tendency of molar cusps to coalesce; and (2) ornamentation of grooves, pits, and ridges on the molars. We argue that the Ptilodontoidea, and less certainly also the Cimolodontidae and Boffiidae, might have originated from amongthe plagiaulacid line, a possible intermediate link being the Paracimexomys group. The remaining Cimolodonta might have originated from unknown members of the Paracimexomys group with separated molar cusps and smooth enamel. The origin of two types of prismatic enamel and a relationship between them are stumbling blocks in understanding the origin of the Cimolodonta; we conclude that microprismatic enamel made its appearance only once. Revised diagnoses of high-rank multituberculate taxa, including lists of all known genera, are given.