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.     

Unique inhibitory cascade pattern of molars in canids contributing to their potential to evolutionary plasticity of diet

Developmental origins that guide the evolution of dental morphology and dental formulae are fundamental subjects in mammalian evolution. In a previous study, a developmental model termed the inhibitory cascade model was established. This model could explain variations in relative molar sizes and loss of the lower third molars, which sometimes reflect diet, in murine rodents and other mammals. Here, I investigated the pattern of relative molar sizes (inhibitory cascade pattern) in canids, a taxon exhibiting a wide range of dietary habits. I found that interspecific variation in canid molars suggests a unique inhibitory cascade pattern that differs from that in murine rodents and other previously reported mammals, and that this variation reflects dietary habits. This unique variability in molars was also observed in individual variation in canid species. According to these observations, canid species have greater variability in the relative sizes of first molars (carnassials), which are functionally important for dietary adaptation in the Carnivora. In conclusion, an inhibitory cascade that differs from that in murine rodents and other mammals may have contributed to diverse dietary patterns and to their parallel evolution in canids.     

Evolutionary and Biological Implications of Dental Mesial Drift in Rodents: The Case of the Ctenodactylidae (Rodentia,Mammalia)

Dental characters are importantly used for reconstructing the evolutionary history of mammals, because teeth represent the most abundant material available for the fossil species. However, the characteristics of dental renewal are presently poorly used, probably because dental formulae are frequently not properly established, whereas they could be of high interest for evolutionary and developmental issues. One of the oldest rodent families, the Ctenodactylidae, is intriguing in having longstanding disputed dental formulae. Here, we investigated 70 skulls among all extant ctenodactylid genera (Ctenodactylus, Felovia, Massoutiera and Pectinator) by using X-ray conventional and synchrotron microtomography in order to solve and discuss these dental issues. Our study clearly indicates that Massoutiera, Felovia and Ctenodactylus differ from Pectinator not only by a more derived dentition, but also by a more derived eruptive sequence. In addition to molars, their dentition only includes the fourth deciduous premolars, and no longer bears permanent premolars, conversely to Pectinator. Moreover, we found that these premolars are lost during adulthood, because of mesial drift of molars. Mesial drift is a striking mechanism involving migration of teeth allowed by both bone remodeling and dental resorption. This dental innovation is to date poorly known in rodents, since it is only the second report described. Interestingly, we noted that dental drift in rodents is always associated with high-crowned teeth favoring molar size enlargement. It can thus represent another adaptation to withstand high wear, inasmuch as these rodents inhabit desert environments where dust is abundant. A more accurate study of mesial drift in rodents would be very promising from evolutionary, biological and orthodontic points of view.     

Phylogenetic memory of developing mammalian dentition

Structures suppressed during evolution can be retraced due to atavisms and vestiges. Atavism is an exceptional emergence of an ancestral form in a living individual. In contrast, ancestral vestige regularly occurs in all members of an actual species. We surveyed data about the vestigial and atavistic teeth in mammals, updated them by recent findings in mouse and human embryos, and discussed their ontogenetic and evolutionary implications. In the mouse incisor and diastema regions, dental placodes are transiently distinct being morphologically similar to the early tooth primordia in reptiles. Two large vestigial buds emerge in front of the prospective first molar and presumably correspond to the premolars eliminated during mouse evolution. The incorporation of the posterior premolar vestige into the lower first molar illustrates the putative mechanism of evolutionary disappearance of the last premolar in the mice. In mutant mice, devious development of the ancestral tooth primordia might lead to their revivification and origin of atavistic supernumerary teeth. Similarity in the developmental schedule between three molars in mice and the respective third and fourth deciduous premolar and the first molar in humans raises a question about putative homology of these teeth. The complex patterning of the vestibular and dental epithelium in human embryos is reminiscent of the pattern of "Zahnreihen" in lower vertebrates. A hypothesis was presented about the developmental relationship between the structures at the external aspect of the dentition in mammals (oral vestibule, pre-lacteal teeth, paramolar cusps/teeth), the tooth glands in reptiles, and the earliest teeth in lower vertebrates.     

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.     

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.     

记中国首次发现的"真古兽类" (eupantotherian)化石

记述了在中国首次发现的采自辽宁黑山县八道壕矿区早白垩世晚期沙海组的一件“eu- pantotherian”(“真古兽类”)下颌骨化石。标本保存了最后两个前臼齿和4个臼齿,它以抬高的下颌角突,半臼齿化的最后一枚前臼齿,臼齿上面积增大但未发育成完整盆形的跟座,尚未形成的facet-5,及加长的最后臼齿等特点有别于所有已知的“eupantotherian”和具有雏形磨楔式臼齿的Kielantherium,被命名为一新属新种,Mozomus shikamai gen.et sp.nov.(鹿间明镇古兽),并由它而创建了一新科,Mozomuridae fam.nov. “Eupantotherian”是早期哺乳动物演化中的一个重要环节,是从无跟座的对齿兽(sym- metrodont)到具有磨楔式(tribosphenic)臼齿兽类的中间类型。早期兽类进化的成功模式是发育成具有磨楔式的臼齿,即在上臼齿上发育出原尖,而下臼齿的跟座形成由3个齿尖围成的盆状。这种结构扩大了牙齿的面积,使咀嚼切割能力更趋完善,今天的有袋类和真兽类均是如此。但在哺乳动物系统发育史上,“eupantotherian”类的化石发现不多,这给探讨具有磨楔式臼齿构造的两大门类(后兽类和真兽类)的起源带来不少困难和疑惑。而传统上的真古兽类形态分异又很大,并不是一个单系类群。其中有跟座发育较好者,如peramurans有可能更接近具有磨楔式臼齿兽类的基部位置,本文记述的Mozomus shikamai也应属于这一类型。具有雏形的被认为处于基干上的磨楔式臼齿类化石,迄今只有两种,即发现在英国早白垩世地层中的滨齿兽(Aegialodon)和蒙古早白垩世晚期Hoobor层的Kielantherium,前者仅有一颗下臼齿,后者由一枚下臼齿和一具有4颗臼齿的下牙床为代表。两种化石在分类上被归入单一的滨齿兽目(Order Aegialodontia Butler,1978),视为Boreosphenidans的基干(stem)。本文记述的Mozomus,其时代与Kielantherium的大体相当,在大小、齿式及臼齿形态上与后者也多有相近之处,但前者以其臼齿的facet-5尚未出现和跟盆发育不全等特点表明它较Kielantherium更为原始,不具备磨楔式臼齿的模式,因之不能归人Aegialodontia,而只能纳入”eupantotheri- ans”。但在后一类的组合中,Mozomus以它半臼齿化的最后前臼齿和面积增大但未发育成盆形的跟座等特点,又是组合中相当进步的类型。无论如何Mozomus的发现是在为数极少的向磨楔式臼齿模式进化的中间环节上增添了一件重要的化石标本,也增加了不少新的信息。它必会引起学者对这一进化过程的更加深入的反思和新的启示。     

A new spalacolestine mammal from the Early Cretaceous Jehol Biota and implications for the morphology,phylogeny, and palaeobiology of Laurasian ‘symmetrodontans’

‘Symmetrodontans’ are extinct mammals characterized by having a reversed‐triangle molar pattern in which three main cusps define a triangular molar crown. This dental morpholgy has been regarded as being intermediate between the ‘triconodont’ tooth and the tribosphenic pattern characterizing therians; it is a key feature in taxonomy of Mesozoic mammals and one to understand mammalian evolution and palaeobiology. Here we report a new genus and species of ‘symmetrodontan’ mammal, Lactodens sheni, from the Early Cretaceous Jehol Biota, represented by a partial skeleton with dentary and upper and lower teeth with dental morphologies well‐preserved. The new species has a dental formula of three upper incisors, one canine, three premolars, and six molars/three lower incisors, one canine, five premolars and six lower molars, double‐rooted canines, extremely low‐crowned and transversely thin premolars, and acute angled molars. The dental morphologies of molars and peculiar deciduous premolars are similar to those of Spalacolestes from North America. The associated upper and lower dentitions from one individual animal helped to clarify tooth identification of some spalacotheriids represented only by fragmentary material. Phylogenetic analyses indicate a close relationship of the new species to North American spalacolestines and faunal interchanges between Eurasia and North America, thus supporting the notion that small‐bodied spalacotheriids were diverse and had a pan‐Laurasian distribution during the Early Cretaceous. Absence of the Meckelian groove suggests acquisition of the definitive mammalian middle ear in spalacolestines, and deciduous canines and premolars in the slim and extremely long dentary imply a faunivorous diet.     

The disposition of the apices of the roots of multirooted teeth in the alveolar processes of the upper jaw]

Specific features of the disposition of root apexes of premolars and molars in the alveolar process of the upper jaw were studied in 148 skulls and 67 tooth-jaw cuts. The least thickness of the alveolar process was found to be at the level of the first premolar (10,41 +/- 0,33 mm), and the greatest--at the level of the first molar (14,075 +/- +/- 0,456 mm). Strong correlation of the alveolar process was established between the levels of the premolars and the last two molars. The distance from the apexes of cheek roots of the first molar to the bottom of the maxillary sinus is 2,05--2,92 mm, and the palatal roots -- 2,02 mm. On roentgenograms the molar roots are often projected on the sinus bottom.     

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.     

Mandibular premolar and second molar root morphological variation in modern humans: What root number can tell us about tooth morphogenesis

This investigation of modern human mandibular premolar root variation tests the hypothesis that population-specific mandibular single-rooted premolar root size can predict a predisposition to root morphological complexity. Mandibular postcanines were examined and quantified from dental radiographs in a globally spread sample of 1,615 modern humans. Multirooted premolars and a fused molar root phenotype were investigated as probes into greater than, and less than, the normative root number. Twelve questions were addressed concerning root structure of mandibular premolars and second molars. A direct correlation was found between single-rooted mandibular premolar size and the predisposition to multirootedness. This correlation infers the following: 1) that postcanine primordia size during root formation predisposes to the development of more, or less, than the normative postcanine root number; and 2) that the epigenetic effect of tooth primordium size per se influences the induction of interradicular processes, which divides the root during its development. This simple developmental model helps explain the following observations: 1) population-specific variation in postcanine root number; 2) sexual dimorphism for multirooted mandibular premolar prevalence; 3) why microdont teeth are single-rooted; 4) the hierarchy of developmental canalization of interradicular processes; 5) megadont-hominin to late-hominin mandibular premolar root number transition; and 6) the fluctuation of mandibular premolar root number in primate evolutionary history.     

3D Mapping of Safe and Danger Zones in the Maxilla and Mandible for the Placement of Intermaxillary Fixation Screws

Intermaxillary (IMF) screws feature several advantages over other devices used for intermaxillary fixation, but using cone beam computed tomography (CBCT) scans to determine the safe and danger zones to place these devices for all patients can be expensive. This study aimed to determine the optimal interradicular and buccopalatal/buccolingual spaces for IMF screw placement in the maxilla and mandible. The CBCT volumetric data of 193 patients was used to generate transaxial slices between the second molar on the right to the second molar on the left in both arches. The mean interradicular and buccopalatal/buccolingual distances and standard deviation values were obtained at heights of 2, 5, 8 and 11 mm from the alveolar bone crest. An IMF screw with a diameter of 1.0 mm and length of 7 mm can be placed distal to the canines (2 - 11 mm from the alveolar crest) and less than 8 mm between the molars in the maxilla. In the mandible, the safest position is distal to the first premolar (more than 5 mm) and distal to the second premolar (more than 2 mm). There was a significant difference (p<0.05) between the right and left quadrants. The colour coding 3D template showed the safe and danger zones based on the mesiodistal, buccopalatal and buccolingual distances in the maxilla and mandible.The safest sites for IMF screw insertion in the maxilla were between the canines and first premolars and between the first and second molars. In the mandible, the safest sites were between the first and second premolars and between the second premolar and first molar. However, the IMF screw should not exceed 1.0 mm in diameter and 7 mm in length.     

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.     

陕西蓝田灞河组的小羚羊(Dorcadoryx Teilhard et Trassaert, 1938, Bovidae)

描述了牛科小羚羊(Dorcadorys Teilhard et Trassaert,1938)一新种:D.orientalis sp. nov.。材料发现于陕西蓝田Loc.6,晚中新世灞河组中部。它和属型种Dorcadoryx triquetricor- nis的主要不同在于它的角心长,向上分散度大,角后窝深而狭长,眼眶向头骨两侧突出不明显,下颌骨水平支低而纤细,颊齿窄和前臼齿列相对长等。Dorcadoryx是中国北方和蒙古地区晚中新世至早上新世的特有类型之一。我国山西晚中新世的?Tragoreas(Bohlin,1935)、蒙古新近纪晚期的Tragoreas sp.2和sp.3以及Olonbulukia(?)sp.(Dmitrieva,1977)可能是它的成员。经修订,属的主要特征是个体小至中等大小,头骨狭长,角基之间的额面不向上隆起,头骨弯曲发生在额顶骨之间,眶上孔小,眶前窝大,颅顶部狭长,基枕部具中沟,前结节不明显,角心位于眼眶之上、不旋转,内外侧扁,分散度不大,稍向后弯曲,角柄长,角后窝存在, 前臼齿列退化以及p4的下后尖和下前尖相连等。它可能属于山羊亚科,包含4—5种。新种可能是其中的一个早期类型。     

Interproximal wear facets and tooth associations in the Paşalar hominoid sample

Interproximal wear facets were examined on hominoid teeth from the middle Miocene site at Pa?alar, Turkey. The aim was to find matches between adjacent premolar and molar teeth from single individuals that were collected in the field as isolated teeth and use them to reconstruct tooth rows. These were then used to investigate: (1) the wear gradient on the molar teeth; (2) the dispersal of teeth from single mandibles and maxillae; (3) the size ratios among the molars; and (4) the number of individuals represented by the hominoid sample. Facets were scored for size and shape and were assessed visually using photographs and superimposed outline drawings on acetate transparencies. Out of a sample of approximately 1,500 teeth collected between 1983 and 1996, 532 molars and 258 premolars produced apparent matches making up 160 tooth rows. These were then examined rigorously for morphological consistency and state of wear, and, employing the criterion that only the most unequivocal associations should be used, the final number was reduced to 48 tooth rows-31 mandibular and 17 maxillary. The tooth associations represent a minimum of 21 individuals and probably as many as 34. Molar wear was rapid, with M1s having almost twice as much wear as M3s, as measured by a wear-gradient index. The M2s are intermediate but generally closer to M1s in degree of wear, as are P4s. This wear pattern suggests either delayed eruption of M3s or extremely abrasive diets causing rapid, heavy wear. There is some indication that the wear patterns in Griphopithecus alpani and Kenyapithecus kizili are different, with the latter perhaps having a lower wear gradient, but the K. kizili sample is very small. In both species, the M2 is the largest molar and the M1 is the smallest. Separation of individual teeth in the 48 tooth associations varied from widely separated-up to 8.5m apart-to within a few centimeters of each other. One tooth row (D922) was found with the teeth in contact but the maxillary bone had dissolved away. Two dispersal mechanisms have been identified from earlier taphonomic work: transport of disarticulated elements to the fossil site and reworking of sediments by spring action.     

Developmental constraints revealed by co-variation within and among molar rows in two murine rodents

SUMMARY Morphological integration corresponds to interdependency between characters that can arise from several causes. Proximal causes of integration include that different phenotypic features may share common genetic sets and/or interact during their development. Ultimate causes may be the prolonged effect of selection favoring integration of functionally interacting characters, achieved by the molding of these proximal causes. Strong and direct interactions among successive teeth of a molar row are predicted by genetic and developmental evidences. Functional constraints related to occlusion, however, should have selected more strongly for a morphological integration of occluding teeth and a corresponding evolution of the underlying developmental and genetic pathways. To investigate how these predictions match the patterns of phenotypic integration, we studied the co‐variation among the six molars of the murine molar row, focusing on two populations of house mice (Mus musculus domesticus) and wood mice (Apodemus sylvaticus). The size and shape of the three upper and lower molars were quantified and compared. Our results evidenced similar patterns in both species, size being more integrated than shape among all the teeth, and both size and shape co‐varying strongly between adjacent teeth, but also between occluding teeth. Strong co‐variation within each molar row is in agreement with developmental models showing a cascade influence of the first molar on the subsequent molars. In contrast, the strong co‐variation between molars of the occluding tooth rows confirms that functional constraints molded patterns of integration and probably the underlying developmental pathways despite the low level of direct developmental interactions occurring among molar rows. These patterns of co‐variation are furthermore conserved between the house mouse and the wood mouse that diverged >10 Ma, suggesting that they may constitute long‐running constraints to the diversification of the murine rodent dentition.     

Report of inaccuracies in the study of the dentition of mammals

After discussing the existing view points and on the base of author's own studies, the designation of premolars and molars is proposed in the deciduous and permanent dentition of placental mammals, corresponding to the function, form, and structure of the teeth. With the admittance of the molars in the deciduous dentition of mammals, the discrepancy between milk dentition of mammals and humans is eliminated. In this way, the molars of deciduous dentition appeared to be the precursors of premolars and molars of permanent dentition. The separate groups of teeth are differentiated depending on the way of life and kind of consumed food, according to position and function of the teeth in the dentition. Lacerating teeth, premolars are formed in carnivora mammals for tearing off the food, and in case of intensive masticatory function of herbivora and omnivora, very likely, from the same germs of premolars, the molars are formed.     

Stable carbon and oxygen isotopes in human tooth enamel: Identifying breastfeeding and weaning in prehistory

This paper investigates the utility of stable carbon and oxygen isotopes in human dental enamel to reveal patterns of breastfeeding and weaning in prehistory. Enamel preserves a record of childhood diet that can be studied in adult skeletons. Comparing different teeth, we used δ¹³C to document the introduction of solid foods to infant diets and δ¹⁸O to monitor the decline of breastfeeding. We report enamel carbonate δ¹³C and δ¹⁸O of 33 first molars, 35 premolars, and 25 third molars from 35 burials from Kaminaljuyú, an early state in the valley of Guatemala. The skeletons span from Middle Preclassic through Late Postclassic occupations, ca. 700 B.C. to 1500 A.D. Sections of enamel were removed from each tooth spanning from the cusp to the cemento-enamel junction. Stable isotope ratios were measured on CO₂ liberated by reaction of enamel with H₃PO₄ in an automated carbonate system attached to a VG Optima mass spectrometer. Within a skeleton, teeth developing at older ages are more enriched in ¹³C and more depleted in ¹⁸O than teeth developing at younger ages. Premolars average 0.5% higher in δ¹³C than first molars from the same skeleton (P = 0.0001), but third molars are not significantly enriched over premolars. The shift from first molars to premolars may be due to the shift to solid foods from lipid-rich milk. After 2 years, when premolars begin to mineralize, the δ¹³C in childhood diets did not change systematically. First molars and premolars are similar in δ¹⁸O, but third molars average 0.7% lower than first molars (P = 0.0001) and 0.5% lower than premolars (P = 0.0003). First molar and premolar δ¹⁸O is heavier, because breast milk is more enriched in ¹⁸O than is drinking water. Hence, many children continued to nurse during the period of premolar formation. Together, these results indicate that Kaminaljuyú children had begun to eat solid maize foods before the age of 2 years but continued to drink breast milk until much later. Am J Phys Anthropol 106:1–18, 1998. © 1998 Wiley-Liss, Inc.     

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.     

Occlusal Pattern in Paulchoffatiid Multituberculates and the Evolution of Cusp Morphology in Mammaliamorphs with Rodent-like Dentitions

Multituberculates developed a very complex masticatory apparatus during their long evolutionary history from the Jurassic to the Paleogene. Besides their rodent-like elongated incisors and diastemata, Cenozoic cimolodont Multituberculata display masticatory movements involving two distinct cycles in the mastication. An orthal slicing-crushing cycle associated with an enlarged lower fourth premolar precedes a palinal grinding cycle linked to upper molars with three longitudinal rows of cusps. With their plesiomorphic lower premolars and upper molars, the Late Jurassic/Early Cretaceous multituberculate family Paulchoffatiidae can provide the key for the understanding of the origin of the complex mastication of the Cimolodonta. Using for the first time propagation phase contrast Synchrotron X-Ray microtomography to perform both microwear and topographic analyses in order to characterize the mastication of Paulchoffatiidae, we digitized dental material from the Late Jurassic of the Guimarota Coal Mine (Leiria, Portugal) at the European Synchrotron Facility (Grenoble, France). Mastication in Paulchoffatiidae is characterized by a palinal grinding cycle. In contrast to Cimolodonta, no evidence of an orthal slicing-crushing cycle has been observed: the lower premolars mainly have a grinding function like the molars as they do exhibit buccal attrition facets bearing longitudinal striations. Nevertheless, the slightly oblique striations observed on the mesial part of the paulchoffatiid lower premolars possibly presage the orthal phase of the Cimolodonta. Our topographic analysis indicates that a strong relationship between individual cusp shape and direction of chewing is emphasized in rodents and rodent-like Mammaliamorpha such as Cimolodonta and Tritylodonta. Surprisingly, this relationship is not evident in Paulchoffatiidae. This unexpected result can be explained by the non-involvement in the attrition of many premolar cusps in Paulchoffatiidae, as indicated by our microwear analysis. The stronger the attrition, the more the direction of the masticatory movements influences the cusp morphology in Mammaliamorpha.