Tooth Eruption Sequences in Cervids and the Effect of Morphology,Life History,and Phylogeny

Tooth eruption sequences vary in a non-random way among mammalian species. Several variables have been linked to this, including tooth and jaw shape, adaptations to diet, and food processing. Likewise, changes in eruption patterns correlate with the speed of postnatal growth in some groups, the Schultz’s Rule pattern. Here, the eruption pattern of the permanent dentition in lower jaws from different cervid species have been investigated to discern the effect of these factors and phylogeny as well as to reconstruct the ancestral tooth eruption sequence of cervids. In ruminants, the different patterns of emergence of permanent teeth seem to be best explained by phylogeny. The degree of hypsodonty, age of first molar eruption, and life history parameters such as longevity and age of female sexual maturity do not explain the observed sequential differences in eruption patterns. The Parsimov-based analysis for the ancestral state resulted in a tooth eruption sequence of m1 – m2 – i1 – i2 – i3 – c – m3 – (ppp) for Cervidae; a pattern recorded in Odocoileus, Capreolus, and Hydropotes. The eruption pattern of Caenomeryx filholi, from the Oligocene of Gaimersheim, is identical to the result of the Parsimov-based analysis except for the presence of a first premolar, a tooth lost in cervids.     

New remains of Hyaenodontidae (Creodonta,Mammalia) from the Oligocene of Central Mongolia

New specimens of hyaenodontid creodonts are described which were unearthed in Mongolia by the Austrian–Mongolian Paleontological expeditions. Five taxa are identified: Hyaenodon eminus, Hyaenodon pervagus, Hyaenodon cf. incertus, Hyaenodon cf. mongoliensis, and cf. Hyaenodon gigas. Hitherto unknown morphological details are reported for most of these species. The implications to systematic of the species and the genus Hyaenodon are discussed. These Hyaenodon remains suggest a separate evolutionary lineage, with different species in Europe and Asia, even if the genus is also known from contemporary faunas of Europe and North America.     

Amyloidity is not diagnostic for species in the Mycena pearsoniana complex (Mycena sectio Calodontes)

In Mycena sectio Calodontes with otherwise amyloid spores, the inamyloid spores of Mycena pearsoniana Dennis ex Singer were a distinguishing feature for this species and its subsection Violacella. Although the original concept of this species was European, Singer chose to typify it with material collected in Mexico. The name has since been applied to all European collections with inamyloid spores and decurrent lamellae. Our phylogenetic analysis of 91 ITS sequences from European, North and South American Calodontes collections shows that European collections identified as M. pearsoniana fall into two well-supported sibling clades together with both inamyloid and weakly amyloid North American collections. Since the holotype of M. pearsoniana is in an advanced state of decay, we have selected an epitype from a North American locality with a climate comparable to the Mexican type locality. Our results show weakly and inamyloid spore reactions to be homoplastic in Calodontes, and furthermore that spores of M. pearsoniana can show either amyloid or inamyloid reactions interchangeably. This raises doubt about the taxonomic value of this trait in Mycena systematics.     

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.     

New material of Diacodexis (Mammalia,Artiodactyla) from the early Eocene of Southern Europe

Diacodexeidae are the first representatives of Artiodactyla in the fossil record. Their first occurrence is at the very base of the Ypresian (earliest Eocene, 56.0 Ma) with Diacodexis, a genus well diversified during the early Eocene in Europe, especially during the MP7–MP8 + 9 interval. However, most of European species are documented by scarce material, retrieved from single localities. In this work, we describe new Diacodexis material from ～MP7 and ～MP8 + 9 localities of Southern Europe, including material of D. antunesi from Silveirinha, considered as the most primitive European Diacodexis species, and material from three localities from Southern France (Fordones, Palette, and La Borie). The new material documents Diacodexis premolar morphology and deciduous dentition which bear potentially important phylogenetic information, as well as astragali, including a specimen from Silveirinha that constitutes the earliest occurrence of an astragalus of the genus Diacodexis in the European fossil record. Investigation of the enamel microstructure reveals that early European species had a simple enamel pattern with one-layered Schmelzmuster composed of ‘basic’ radial enamel only, instead of the two-layered Schmelzmuster (thin radial enamel + thick layer of Hunter-Schreger bands) observed on North American species and so far considered to represent the primitive condition within Artiodactyla. In accordance with previous studies, our observations highlight that Diacodexis gigasei from Belgium is morphologically closer to the North American species D. ilicis than to D. antunesi from Portugal. The latter species, together with D. aff. antunesi from Fordones, appears to be morphologically closer to the Asiatic taxa D. indicus and D. pakistanensis. Finally, we found numerous similarities between D. cf. gigasei from Palette and D. gigasei, a result that challenges the intra-European provincialism that characterizes the earliest Ypresian. Diacodexis gigasei could be one of the rare species shared by the northwestern and southwestern European bioprovinces.     

Fabrosauridae,the basal family of ornithischian dinosaurs (Reptilia: Ornithopoda)

In fabrosaurids the upper jaw is flat and the lower jaw is slender so the ’cheek’ teeth are marginal and not inset as is the case in all other ornithischian dinosaurs. The ’cheek’ teeth of fabrosaurids have anteroposteriorly expanded crowns but lack wear surfaces formed by tooth to tooth contact. Two genera are recognized from the Triassic-Jurassic boundary of Lesotho with good material previously referred toFabrosaurus as a new genus that represents the most conservative ornithopod described to date. The anatomy ofNanosaurus (Upper Jurassic, U.S.A.) andEchinodon (Jurassic-Cretaceous boundary, England) is redescribed; in both genera the tooth bearing bone of the lower jaw is deepened posteriorly and inEchinodon there is a true canine tooth in the upper jaw.     

Aspects of dental development in the orangutan prior to eruption of the permanent dentition

In spite of a resurgence of interest in the interpretation of the sequences of dental development and eruption in various Plio-Pleistocene hominoids as being either “modern human” or “ape-like,” the body of comparative material on the extant hominoids remains deficient in critical areas. In concert with recent attempts to rectify this situation, we present the results of our studies on dental morphogenesis in the orangutan. We have focused on the growth and eruption of the deciduous dentition as well as early stages of permanent tooth formation and have found that 1) many permanent teeth develop earlier than was thought, 2) differences exist between development in the upper and in the lower jaw, and 3) states of tooth formation can vary significantly among individuals of similar chronological age or tooth eruption status.     

Behavioral correlates of tooth eruption in Madagascar lemurs

Observations on the sequence and timing of gingival tooth eruption are reported for six species of Madagascar lemurs. Complete sequences of eruption were obtained for the deciduous dentition, and partial to complete sequences were recorded for the permanent dentition. In Cheirogaleus medius and in four species of the genus Lemur, the deciduous teeth erupt in front-to-back sequence, with the toothcomb emerging near birth as an integrated complex. In Propithecus verreauxi the same pattern is exhibited, but the small peglike lower canine and dp₃ erupt last. Eruption of the permanent dentition in Lemur species takes place in two distinct stages. In the first stage the upper incisors, toothcomb, and first two molars penetrate the gingiva. After an interval of 3 to 4 months, the remaining permanent teeth erupt. Deciduous premolars erupt when young animals are being weaned. The eruption of the deciduous toothcomb appears unrelated to feeding or grooming behavior. In L. catta and L. fulvus, the first stage of permanent tooth eruption occurs at approximately 6 months of age, when the growth rate slows down and (in wild populations) the rainy season is ending. This suggests that eruption of the anterior molars is timed to coincide with a shift from a more frugivorous to a more folivorous dietary regime, which occurs during the dry season. No further tooth eruption occurs until approximately 1 year of age, when the growth rate increases and the rainy season returns for wild populations. Thus, the second wave of permanent tooth eruption in these species again appears linked to changing climatic conditions which lead to a shift in dietary preferences.     

Dental Eruption Series and Replacement Pattern in Miocene <Emphasis Type="Italic">Prosantorhinus</Emphasis> (Rhinocerotidae) as Revealed by Macroscopy and X-ray: Implications for Ontogeny and Mortality Profile

Fossil evidence of complete sequences of dental ontogeny in extinct mammals is rare but contains valuable information on the animal’s physiology, life history, and individual age. Here, we analyzed an exceptionally high number of juvenile dentaries at different developmental stages including highly fragile tooth germs of the extinct rhinoceros Prosantorhinus germanicus from the Miocene fossil lagerstätte Sandelzhausen in Germany. We used dental wear stages, eruption stages, and tooth germ development in order to reconstruct the tooth replacement pattern for P. germanicus. The results allow for the distinction of 11 dental eruption stages and document a tooth eruption sequence of (d2, d3), (d1, d4), m1, m2, p2, p3, p4, m3; a pattern identical to that reported for the extant African rhinoceros, Diceros bicornis. Moreover, our findings indicate that P. germanicus falls into the life history category of slow-growing, long-living mammals. The dental eruption stages of the fossil rhinoceros were correlated with data of living rhinoceroses in order to gain insight into the age-at-death distribution of P. germanicus at Sandelzhausen. The juvenile mortality profile of P. germanicus shows a trend of selective mortality at an inferred age range of about 3 months to 3 years. As this age range represents a life phase of increased natural risk of mortality, our findings indicate a gradual accumulation of corpses (attritional fossil assemblage). This result supports the interpretation of a taphocenosis found at the Sandelzhausen fossil site.     

The eruption times of permanent teeth in boys and girls in the Stormarn District, Schleswig-Holstein (Germany)

In a longitudinal study in two small towns in southern Schleswig-Holstein (Ammersbek and Ahrensburg, District Stormarn; 9155 inhabitants) we investigated 2832 oral findings of 1396 patients (711 males, 685 females). The minimum age was 1.51 years, and the maximum age was 25.50 years. The dental findings were collected over a period of about 20 years (1982-2002). The oral findings per child were assessed between one and eight times. The eruption times of teeth in females are earlier than those for the same teeth in males. Further, the permanent dentition in females is completed earlier than in males. In both sexes the tooth eruption occurs symmetrically in both jaws. The comparison of both jaws revealed a slightly advanced eruption of the lower jaw teeth in both sexes. There is a noteworthy change in the eruption sequence of the teeth. In contrast to other reports we observed that the eruption of the canine proceeds the eruption of the second molar. We found no acceleration of the dentition when compared with other reports and could confirm the rules of tooth eruption in man. Conclusion: Oral examination of teeth is a simple tool to calculate tooth eruption intervals. This first investigation on a population of Schleswig-Holstein revealed a change in the eruption sequence of permanent teeth. These findings are relevant for dental treatment planning and should be reconfirmed at certain intervals.     

Occlusal and morphogenetic field evolution in the dentition of European Nyctitheriidae (Euarchonta,Mammalia)

Abstract

The nyctithere genera Saturninia, Cryptotopos and members of the subfamily Amphidozotheriinae are well-represented by dental remains in the European later Eocene. Their molar occlusal relations are described in detail, demonstrating a diversity of adaptations to insectivory, including dilambdodonty, minor trends in zalambdodonty, development of a large talon shelf in upper first and second molars and a step in the trigon-trigonid shearing surface that provides an extra facet in buccal phase. Minor lingual phase wear is recognized for the first time in the family, but only in relatively worn teeth. Nyctithere molars differ from those of most insectivorous mammals today in having well-developed paraconules and metaconules on upper molars, which in most cases lack a marked bucco-lingual tilt, associated with a more transverse jaw action. Amphidozotherium, however, shows tilting and a basally shifted M¹ hypocone associated with M₁ talonid exodaenodonty and a minor trend in zalambdodonty. Nyctitheres primitively have three deeply notched lobate lower incisors and a large but procumbent premolariform lower canine. Amphidozotheriines have modified their I₃ into a premolariform tooth, by shifting the premolarization field mesially. Amphidozotherium has also shifted this field distally, reducing P_2–3 in size and their roots from two to one.     

A new species of Pliopithecus from the Indian S̀ivaliks

This paper gives a detailed account of an isolated upper third molar of a gibbon-like primate, the first of its kind ever found from the S?ivalik hills. The tooth comes from the Nagri beds comparable to the Middle Miocene Formations. This tooth has been assigned to a new species of Pliopithecus, namely P. krishnaii, on the basis of morphological and metrical comparisons with the fossil gibbon-like primates and living gibbons.     

The “miacids” (Carnivoraformes,Mammalia) from the Early Eocene locality of Le Quesnoy (MP7, France); first occurrence of Vassacyon in Europe

We describe here “miacid” taxa from the Early Eocene Paris Basin locality of Le Quesnoy (Oise, France). We describe the new species Vassacyon taxidiotis, the first European record of this genus. The other “miacids” identified from Le Quesnoy are Miacis latouri and Gracilocyon solei. The P4 of G. solei is described here for the first time. Its morphology (e.g., wide protocone, short postmetacrista) supports a close relationship with Miacis rundlei from Abbey Wood (MP8 + 9, England). The latter species is therefore classified as Gracilocyon rundlei. Three new tooth positions are known for Miacis latouri: P4, p4 and m2. They support its reference to Miacis. These specimens imply that the European species is more basal than the North American species. The fauna from Le Quesnoy shares with Dormaal the presence of Miacis latouri and Gracilocyon solei, but the “miacid” fauna from Le Quesnoy also contains Vassacyon taxidiotis. The presence in Le Quesnoy of the two former taxa supports a reference to MP7 level of the French locality. The presence of three distinct genera in European localities show that the “Miacidae” were diversified in Europe, as previously observed in North America. The genera Gracilocyon, Miacis, and Vassacyon probably dispersed from Europe to North America during the Palaeocene/Eocene boundary.     

New dentaries of Chiromyoides (Primatomorpha,Plesiadapidae) and a reassessment of the “mammalian woodpecker” ecological niche

Based on their relatively large, chisel-like incisors and robust dentaries, species of the Paleocene plesiadapid mammal Chiromyoides have been described as potential ecological analogues of either seed-eating rodents or the unusually specialized lemur Daubentonia madagascariensis. Here, we analyze the most complete dentaries of Chiromyoides currently known in order to illuminate jaw form and function in this taxon. Principal Component Analysis shows that Chiromyoides campanicus and Daubentonia are uniquely similar in select dentary proportions when compared with a sample including seven other fossil plesiadapid taxa as well as 22 extant primates, dermopterans and scandentians. Comparative allometric analyses indicate that in both Daubentonia and Chiromyoides, the unique jaw proportions are likely achieved through hypertrophy of masseteric fossa length and dentary depth, rather than simple reduction of tooth row length. Consistent with these dentary features indicative of powerful gnawing, we show that incisor apex morphology became increasingly chisel-like in certain younger species of Chiromyoides. Importantly, slight reduction in molar area relative to jaw length and body mass appears to characterize all species of Chiromyoides in which molar proportions can be estimated. Notably, this pattern occurs in one of the oldest known specimens of Chiromyoides, an edentulous but relatively complete dentary from the middle Tiffanian of Texas, which differs from other Chiromyoides specimens in having a relatively shallower corpus. Taken together, this evidence suggests that Chiromyoides was a Daubentonia-like extractive forager that evolved from taxa whose diets emphasized exudates.     

An onychodont fish (Osteichthyes,Sarcopterygii) from the Early Devonian of China,and the evolution of the Onychodontiformes

Although the superbly preserved specimens of Onychodus jandemarrai have greatly advanced our understanding of the Onychodontiformes, a primitive sarcopterygian group with large parasymphysial tooth whorls, the scarcity of the otoccipital material in the group hampers further morphological comparisons between onychodonts and other sarcopterygian groups. Here we report a new onychodont Qingmenodus yui gen. et sp. nov. from the Early Devonian (Pragian) of South China that comprises well-ossified otoccipital and upper and lower jaw material. As one of the oldest known onychodonts, Qingmenodus shows for the first time the nearly complete structure of the otoccipital in onychodonts and provides an additional basis to address the phylogenetic position of the group. Its elongated otic shelf exhibits the posterior shift of the attachment for the basicranial muscle as in coelacanths and sheds light on the feeding mechanism of onychodonts. Qingmenodus displays a mosaic of primitive and derived onychodont features. The phylogenetic analysis places Qingmenodus immediately basal to the clade comprising Onychodus and Grossius.     

The evolution and paleodiet of the Eurygnathohippus feibeli lineage in Africa

Eurygnathohippus feibeli is a small African species of hipparionine horse that is known from Ethiopia, Kenya, and Libya from the latest Miocene to the early Pliocene. It is characterized by generally primitive and moderately hypsodont maxillary cheek teeth but advanced, elongate distal limb elements. This study focused on developing our understanding of the evolution of paleodiet in the E. feibeli (sensu lato) clade, based on two different methods of analyzing tooth mesowear on upper molars from Lothagam, Kenya, and localities in the Middle Awash, Ethiopia. When tooth cusp shape—as described by two variables for paleodietary reconstruction—was considered, all of the studied samples clustered together with recent species that feed on graze, with the majority clustering together with less extreme grazers like waterbuck and sable antelope and a minority presenting the greatest similarity to extreme grazers such as tsessebe and white rhino. The mesowear method involving a univariate standard reveals scores that compare best with values of recent grazing and some mixed-feeding ungulates, suggesting a similar dietary range within the E. feibeli lineage, and which show a modest trend over time from E. feibeli to E. aff. feibeli towards lower and blunter cusps. While older populations of E. feibeli sensu strictu must have included more browse in their diet, younger populations increasingly preferred graze. The E. feibeli sample may represent an intermediate step within a likely general African hipparionine dietary trend that evolved from more mixed feeding towards extreme grazing through the late Miocene–Pleistocene interval. Some results indicate parallels in the dietary adaptation of North American and African late Miocene to earliest Pliocene equid faunas.     

Morphofunctional Analysis of the Quadrate of Spinosauridae (Dinosauria: Theropoda) and the Presence of Spinosaurus and a Second Spinosaurine Taxon in the Cenomanian of North Africa.

Six quadrate bones, of which two almost certainly come from the Kem Kem beds (Cenomanian, Upper Cretaceous) of south-eastern Morocco, are determined to be from juvenile and adult individuals of Spinosaurinae based on phylogenetic, geometric morphometric, and phylogenetic morphometric analyses. Their morphology indicates two morphotypes evidencing the presence of two spinosaurine taxa ascribed to Spinosaurus aegyptiacus and? Sigilmassasaurus brevicollis in the Cenomanian of North Africa, casting doubt on the accuracy of some recent skeletal reconstructions which may be based on elements from several distinct species. Morphofunctional analysis of the mandibular articulation of the quadrate has shown that the jaw mechanics was peculiar in Spinosauridae. In mature spinosaurids, the posterior parts of the two mandibular rami displaced laterally when the jaw was depressed due to a lateromedially oriented intercondylar sulcus of the quadrate. Such lateral movement of the mandibular ramus was possible due to a movable mandibular symphysis in spinosaurids, allowing the pharynx to be widened. Similar jaw mechanics also occur in some pterosaurs and living pelecanids which are both adapted to capture and swallow large prey items. Spinosauridae, which were engaged, at least partially, in a piscivorous lifestyle, were able to consume large fish and may have occasionally fed on other prey such as pterosaurs and juvenile dinosaurs.     

On the eruption pattern of the permanent incisors and first permanent molars in Paranthropus

It has been claimed recently that Australopithecus exhibited a pattern of permanent tooth eruption like that of extant great apes, whereas a significantly different pattern was shared by Paranthropus and Homo (Dean, 1985). More particularly, each of the four Paranthropus specimens examined in that study was held to show advanced development and eruption of the permanent incisors relative to the first molar. It is demonstrated here that the eruption sequence that was posited for at least one of these four Paranthropus specimens (SK 61) is clearly erroneous, while the developmental/eruption sequences manifested by the other three specimens would appear to be more ambiguous than was claimed. Another juvenile specimen of Paranthropus (KNM-ER 1820) that was not included in Dean's study also does not necessarily support the eruption pattern that was said to characterize that taxon.     

Human skeletal remains from the cave of Hearths,Makapansgat, Northern Transvaal

A detailed study has been made of the oldest South African Upper Pleistocene hominid remains found in Acheulian context in a well-stratified sealed cave deposit, the Cave of Hearths, Makapansgat, Northern Transvaal. Possibly 55,000 years of age, the remains comprise a juvenile right mandibular body with teeth, and part of a right radius. The mandible is highly robust, markedly prognathous, has a slight to moderate bony chin, an appreciable planum alveolare, a low supraspinous foramen, large alveolar part with big tooth roots, parallel upper and lower borders, a superior transverse torus and poorly developed genial apophysis. The teeth are fairly large, narrow and elongate; M₂ is smaller than M₁; both molars have a +5 cusp pattern, and the first molar shows moderate taurodontism. There is good evidence that the jaw shows congenital lack of M₃: after the Chinese Lantian jaw, this is the second oldest hominid mandible and the first African fossil man with this feature. The radius has a relatively large head atop a disproportionately narrow neck; marked angulation of neck on shaft; and a strongly developed bicipital tubercle. The remains show a cluster of features which ally them with African Neandertaloids and earlier hominids of N.W. Africa. These geographically widespread African remains may represent a transitional population between H. erectus and H. sapiens neanderthalensis. This population has been called by Campbell, this author and others H. sapiens rhodesiensis (after the first-discovered specimen from Broken Hill): to this taxon the Cave of Hearths bones are tentatively assigned.