Dental histology of Coelophysis bauri and the evolution of tooth attachment tissues in early dinosaurs

Studies of dinosaur teeth have focused primarily on external crown morphology and thus, use shed or in situ tooth crowns, and are limited to the enamel and dentine dental tissues. As a result, the full suites of periodontal tissues that attach teeth to the jaws remain poorly documented, particularly in early dinosaurs. These tissues are an integral part of the tooth and thus essential to a more complete understanding of dental anatomy, development, and evolution in dinosaurs. To identify the tooth attachment tissues in early dinosaurs, histological thin sections were prepared from the maxilla and dentary of a partial skull of the early theropod Coelophysis bauri from the Upper Triassic (Rhaetian‐ 209–201 Ma) Whitaker Quarry, New Mexico, USA. As one of the phylogenetically and geologically oldest dinosaurs, it is an ideal candidate for examining dental tissues near the base of the dinosaurian clade. The teeth of C. bauri exhibited a fibrous tooth attachment in which the teeth possessed five tissues: enamel, dentine, cementum, periodontal ligament (PDL), and alveolar bone. Our findings, coupled with those of more recent studies of ornithischian teeth, indicate that a tripartite periodontium, similar to that of crocodilians and mammals, is the plesiomorphic condition for dinosaurs. The occurrence of a tripartite periodontium in dinosaurs adds to the growing consensus that the presence of these tissues is the plesiomorphic condition for the major amniote clades. Furthermore, this study establishes the relative timing of tissue development and growth directions of periodontal tissues and provides the first comparative framework for future studies of dinosaur periodontal development, tooth replacement, and histology. J. Morphol. 277:916–924, 2016. © 2016 Wiley Periodicals, Inc.     

Neoselachian sharks from the Callovian–Oxfordian (Jurassic) of Ogrodzieniec,Zawiercie Region,southern Poland

Abstract: Callovian and Oxfordian strata in Ogrodzieniec near Zawiercie, southern Poland, have yielded two shark tooth assemblages that collectively include 14 neoselachian taxa. A previously unrecognised member of the Orectolobiformes, Akaimia altucuspis gen. et sp. nov., is described and characterised by a dentition remarkably similar to modern wobbegong sharks (Orectolobidae) by convergence. The assemblages also include the first anterior teeth ever found of the palaeospinacid ‘Synechodus’prorogatus Kriwet, in addition to teeth from two other palaeospinacids, Sphenodus spp., four different orectolobiforms, two hexanchids and Protospinax spp. These shark tooth assemblages contribute to the poorly known Callovian and Oxfordian neoselachian faunas and indicate that the diversity was higher than previously appreciated, particularly within the Orectolobiformes.     

A new species of Azendohsaurus (Diapsida: Archosauromorpha) from the Triassic Isalo Group of southwestern Madagascar: cranium and mandible

Abstract: Here, we describe a new species of Azendohsaurus from the Middle–Late Triassic of Madagascar, extending the geographical range of a taxon known otherwise only by a single species from Morocco. Although Azendohsaurus has consistently been regarded as an early dinosaur (based on various advanced dental and gnathic features resembling those characterizing certain dinosaur subgroups), the relatively complete skeletal material, now available from Madagascar, argues strongly against its dinosaurian affinities. Rather, the retention of numerous primitive cranial and postcranial features indicates a surprisingly early divergence of Azendohsaurus within Archosauromorpha and an unusual mosaic of characters in this taxon. Features considered diagnostic of Sauropodomorpha thus are inferred to occur homoplastically in at least one clade of nondinosaurian archosauromorphs, indicating a complex evolution and distribution of features traditionally thought to be derived within archosaurs. Azendohsaurus has teeth resembling those of both early sauropodomorph and ornithischian dinosaurs, yet also possesses numerous inarguable basal archosauromorph cranial and postcranial attributes. This highlights the risk of uncritically referring isolated, Middle–Late Triassic (or even later), ‘leaf‐shaped’ teeth with denticles to the Dinosauria. Similarly, the occurrence of such teeth in an early diverging archosauromorph indicates that specializations for herbivory originated more frequently within this clade than conventionally assumed. For example, Azendohsaurus and numerous basal sauropodomorph dinosaur taxa share an array of convergently acquired features associated with herbivory, including tooth denticles, expanded tooth crowns, a downturned dentary and the articular located at the ventral margin of the mandible. Some of these features (denticles, expanded crowns and the ventrally deflected articular) are even more widespread among archosauromorphs, including aetosaurs, silesaurs and ornithischian dinosaurs. A downturned dentary also occurs in Trilophosaurus, a taxon further marked by unique specializations for herbivory, including transversely lophate, tricuspid teeth. An array of features associated with herbivory also occurs in rhynchosaurs and certain crocodilians (e.g. Simosuchus). This distribution suggests that craniodental features associated with herbivory were much more pervasive across the archosauromorph clade than previously recognized, possibly evolving at least six to eight times independently.     

The Late Triassic pseudosuchian Revueltosaurus callenderi and its implications for the diversity of early ornithischian dinosaurs

A new discovery of skeletons of Revueltosaurus callenderi from the Upper Triassic Chinle Formation of Petrified Forest National Park, Arizona clearly shows that Revueltosaurus is not an ornithischian dinosaur as previously supposed. Features such as the presence of a postfrontal, crocodile-normal ankle and paramedian osteoderms with anterior bars place R. callenderi within the Pseudosuchia, closer to crocodylomorphs than to dinosaurs. Therefore, dental characters previously used to place Revueltosaurus within the Ornithischia evolved convergently among other archosaur taxa, and cannot be used to diagnose ornithischian dinosaur teeth. As a result, all other putative North American Late Triassic ornithischians, which are all based exclusively on teeth, are cast into doubt. The only reasonably well-confirmed Late Triassic ornithischians worldwide are Pisanosaurus mertii and an unnamed heterodontosaurid from Argentina. This considerably changes the understanding of early dinosaur diversity, distribution and evolution in the Late Triassic.     

The shark fauna from the Middle Triassic (Anisian) of North-Western Nevada

The shark fauna from the Anisian of Nevada is dominated by durophagous hybodontiforms but also shows an important neoselachian component. Two new species of hybodontiform sharks, Acrodus cuneocostatus and Polyacrodus bucheri , are described in addition to a new neoselachian taxon: Mucrovenator minimus. The enameloid of the teeth of Acrodus and Polyacrodus comprises two layers, an outer compact layer and an inner bundled layer. For the typical three-layered enameloid of neoselachian teeth, we propose to replace the terms parallel-fibred enameloid and tangled-fibred enameloid by the more appropriate parallel-bundled and tangled-bundled enameloid.     

A palaeospinacid shark from the Upper Triassic of south-west England

The fin spines of a new neoselachian shark are described from die Upper Triassic of Aust Cliff, Avon and Holwell, near Frome in Somerset. The spines are assigned to Palaeospinax rhaeticus sp. nov. This record adds to the known diversity of Upper Triassic marine vertebrates from Britain and extends the range of the palaeospinacid sharks into the Rhaetian.     

Chondrichthyan tooth enameloid: past,present, and future

Enameloid is a hard mineralized tissue covering chondrichthyan and actinopterygian teeth. Over the past 40 years, it has been extensively studied in various extinct and extant sharks, leading to the broad use of microstructural characters to differentiate between hybodont and neoselachian teeth. However, the chondrichthyan taxic diversity is disproportionately high compared to the number of taxa explored for enameloid microstructure, and the generalization of these few observations to the whole group is problematic. Indeed, many other groups, in particular modern rays and skates, have been completely overlooked, and almost nothing is known about their tooth histology. Furthermore, the recent discovery of typical neoselachian character in cladodontomorph sharks teeth clearly indicates that we have had an over‐simplified perception of the chondrichthyan enameloid distribution, which put into question the previously proposed evolutive scenarios dealing whith this tissue. We propose a brief historical overview of the study and understanding of chondrichthyan enameloid diversity and briefly discuss preparation issues encountered when dealing with the study of chondrichthyan hypermineralized tissues. Then, the variation of enameloid microstructures encountered in ctenacanthiforms, hybodonts, selachimorphs, and batomorphs is explored, summarized, and discussed. Although the full extent of the diversity and variability of the enameloid microstructure in many of these groups and others remains to be fully determined, we are able to show that most possess a much more complex enameloid microstructure than expected, and propose a revised and more fitting chondrichthyan enameloid terminology, based on the recognition of two main units: an external Single Crystallite Enameloid (SCE) and an internal Bundled Crystallite Enameloid (BCE). Our study reveals new insights in the understanding of character distribution among batomorphs and sets a framework for tackling global chondrichthyan tooth enameloid evolution. © 2015 The Linnean Society of London     

A new heterodontosaurid dinosaur (Reptilia: Ornithischia) from the Upper Triassic Red Beds of Lesotho

A new species of ornithischian dinosaur ( Lycorhinus consors sp. nov.) is established on a skull from the Upper Triassic Red Beds of Lesotho. This ornithischian is assigned to the family Heterodontosauridae of the suborder Ornithopoda. The dinosaurs of the family Heterodonto-sauridae are reviewed: Geranosaurus atavus Broom (1911) is considered a nomendubium and the genus name Heterodontosaurus Crompton & Charig (1962) is held to be a junior synonym for Lycorhinus Haughton (1924).
Functional and palaeoecological implications of the heterodontosaurid dentition are discussed. The pattern of tooth wear may reflect a highly specialized jaw action which involved protraction and retraction of the mandible to produce a grinding effect between upper and lower cheek teeth. Lycorhinus consors is presumed to be a female heterodontosaurid because it differs from all other heterodontosaurids in lacking caniniform tusks. It is suggested that the tusks of heterodontosaurids were functionally analogous to those of tayassuids and tragulids and that they were employed as weapons for intra-specific combat and defence. Dental peculiarities indicate that tooth replacement processes were suppressed in heterodontosaurids; replacement of the teeth seems to have been restricted to a brief period each year (presumably when heterodontosaurids underwent aestivation or hibernation).
A new diagnosis is formulated for the family Heterodontosauridae. The relationships of early ornithopod dinosaurs are briefly reviewed and a new classification is proposed. Ten families of ornithopod dinosaurs are recognized; these are ranked in two grades-one (named Dolichopoda) representing the conservative main stem of the ornithischian phylogenetic tree and the other (named Brachypoda) comprising the several more advanced lines of ornithopod evolution.     

The anatomy of the basal ornithischian dinosaur Eocursor parvus from the lower Elliot Formation (Late Triassic) of South Africa

Ornithischia is a morphologically and taxonomically diverse clade of dinosaurs that originated during the Late Triassic and were the dominant large‐bodied herbivores in many Cretaceous ecosystems. The early evolution of ornithischian dinosaurs is poorly understood, as a result in part of a paucity of fossil specimens, particularly during the Triassic. The most complete Triassic ornithischian dinosaur yet discovered is Eocursor parvus from the lower Elliot Formation (Late Triassic: Norian–Rhaetian) of Free State, South Africa, represented by a partial skull and relatively complete postcranial skeleton. Here, the anatomy of Eocursor is described in detail for the first time, and detailed comparisons are provided to other basal ornithischian taxa. Eocursor is a small‐bodied taxon (approximately 1 m in length) that possesses a plesiomorphic dentition consisting of unworn leaf‐shaped crowns, a proportionally large manus with similarities to heterodontosaurids, a pelvis that contains an intriguing mix of plesiomorphic and derived character states, and elongate distal hindlimbs suggesting well‐developed cursorial ability. The ontogenetic status of the holotype material is uncertain. Eocursor may represent the sister taxon to Genasauria, the clade that includes most of ornithischian diversity, although this phylogenetic position is partially dependent upon the uncertain phylogenetic position of the enigmatic and controversial clade Heterodontosauridae. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 160 , 648–684.     

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.     

The evolution of dinosaur tooth enamel microstructure

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

Lamniform Shark Teeth from the Late Cretaceous of Southernmost South America (Santa Cruz Province,Argentina)

Here we report multiple lamniform shark teeth recovered from fluvial sediments in the (Campanian-Maastrichtian) Cerro Fortaleza Formation, Santa Cruz Province, Argentina. This small tooth assemblage is compared to various lamniform sharks possessing similar dental morphologies, including Archaeolamna, Cretalamna, Dwardius, Dallasiella, and Cretodus. Although the teeth share numerous morphological features with the genus Archaeolamna, including a developed neck that maintains a relatively consistent width along the base of the crown, the small sample size and incomplete nature of these specimens precludes definitive taxonomic assignment. Regardless, the discovery of selachian teeth unique from those previously described for the region broadens the known diversity of Late Cretaceous South American sharks. Additionally, the discovery of the teeth in fluvial sandstone may indicate a euryhaline paleobiology in the lamniform taxon or taxa represented by this tooth assemblage.     

A palaeoequatorial ornithischian and new constraints on early dinosaur diversification

Current characterizations of early dinosaur evolution are incomplete: existing palaeobiological and phylogenetic scenarios are based on a fossil record dominated by saurischians and the implications of the early ornithischian record are often overlooked. Moreover, the timings of deep phylogenetic divergences within Dinosauria are poorly constrained owing to the absence of a rigorous chronostratigraphical framework for key Late Triassic–Early Jurassic localities. A new dinosaur from the earliest Jurassic of the Venezuelan Andes is the first basal ornithischian recovered from terrestrial deposits directly associated with a precise radioisotopic date and the first-named dinosaur from northern South America. It expands the early palaeogeographical range of Ornithischia to palaeoequatorial regions, an area sometimes thought to be devoid of early dinosaur taxa, and offers insights into early dinosaur growth rates, the evolution of sociality and the rapid tempo of the global dinosaur radiation following the end-Triassic mass extinction, helping to underscore the importance of the ornithischian record in broad-scale discussions of early dinosaur history.     

Vertebrate microremains from the Early Cretaceous of southern Tunisia

Microremains of various sharks, actinopterygians and crocodiles have been recovered from two sites in the Douiret Formation and three sites in the Aïn el Guettar Formation in southern Tunisia. The presence of an actinistian is also suggested based on histological study of hemisegments of lepidotrichia. Convergence in dental enameloid microstructure between neoselachian sharks and actinopterygians sharing a tearing dentition is also documented. The vertebrate assemblage of the Douiret Formation suggests a pre-Aptian age for this formation and the presence of Bernissartia in the Aïn el Guettar Formation confirms faunal exchange between Africa and Europa during the Early Cretaceous.     

The homology and phylogeny of chondrichthyan tooth enameloid

A systematic SEM survey of tooth microstructure in (primarily) fossil taxa spanning chondrichthyan phylogeny demonstrates the presence of a superficial cap of single crystallite enameloid (SCE) on the teeth of several basal elasmobranchs, as well as on the tooth plates of Helodus (a basal holocephalan). This suggests that the epithelial-mesenchymal interactions required for the development of enameloid during odontogenesis are plesiomorphic in chondrichthyans, and most likely in toothed gnathostomes, and provides phylogenetic support for the homology of chondrichthyan and actinopterygian enameloid. Along the neoselachian stem, we see a crownward progression, possibly modulated by heterochrony, from a monolayer of SCE lacking microstructural differentiation to the complex triple-layered tooth enameloid fabric of neoselachians. Finally, the occurrence of fully-differentiated neoselachian enameloid microstructure (including compression-resistant tangle fibered enameloid and bending-resistant parallel fibered enameloid) in Chlamydoselachus anguineus, a basal Squalean with teeth that are functionally "cladodont," is evidence that triple-layered enameloid microstructure was a preadaption to the cutting and gouging function of many neoselachian teeth, and thus may have played an integral role in the Mesozoic radiation of the neoselachian crown group.     

Theropod dinosaur teeth from the lowermost Cretaceous Rabekke Formation on Bornholm, Denmark

The dinosaur fauna of the palynologically dated lower Berriasian Skyttegård Member of the Rabekke Formation on the Baltic island of Bornholm, Denmark, is represented by isolated tooth crowns. The assemblage is restricted to small maniraptoran theropods, assigned to the Dromaeosauridae incertae sedis and Maniraptora incertae sedis. The dromaeosaurid teeth are characterized by their labiolingually compressed and distally curved crowns that are each equipped with a lingually flexed mesial carina and a distinctly denticulated distal cutting edge. A morphologically aberrant tooth crown (referred to as Maniraptora incertae sedis) has triangular denticles of uneven width, a feature occasionally found in Upper Cretaceous hesperornithiform toothed diving birds, but also in premaxillary teeth of the velociraptorine Nuthetes from the Lower Cretaceous of England.     

Tooth replacement rates in early chondrichthyans: a qualitative approach

The continuous replacement of teeth throughout their lifetime is a common characteristic of most chondrichthyans. This process was already present in the earliest representatives of the group. It has been well established that different species of extant sharks show rapid tooth replacement rates; however, some authors have suggested that in early chondrichthyans this rate might have been much slower. Here we present a qualitative approach to analyse tooth replacement rates in the Early Devonian shark Leonodus carlsi , the earliest tooth-bearing shark known to date. For this, we have examined 1,103 isolated teeth from Celtiberia, Spain. Our study provides strong evidences of an extremely slow dental replacement in this primitive chondrichthyan based on three independents analyses: (1) statistical analysis of the wear degree, demonstrating that teeth remain functional for a long period of time; (2) analysis of both the histological and the morphological features of the teeth cusps suggests that this chondrichthyan used a maturation process that optimizes its function, thus worn teeth show an efficient working shape that implies their teeth remained functional for a long time after being modelled by use; and (3) estimations of size increments between teeth (Δs) of the same dental family for some recent sharks whose rates of replacement were known prove that Δs is inversely proportional to the rate of replacement ( R ² = 0.8327). The estimated values of tooth replacement rates obtained from Δs for L. carlsi and for some Late Devonian cladoselachian sharks are significatively slower than those observed in current sharks.     

Lower limits of ornithischian dinosaur body size inferred from a new Upper Jurassic heterodontosaurid from North America

The extremes of dinosaur body size have long fascinated scientists. The smallest (<1 m length) known dinosaurs are carnivorous saurischian theropods, and similarly diminutive herbivorous or omnivorous ornithischians (the other major group of dinosaurs) are unknown. We report a new ornithischian dinosaur, Fruitadens haagarorum, from the Late Jurassic of western North America that rivals the smallest theropods in size. The largest specimens of Fruitadens represent young adults in their fifth year of development and are estimated at just 65–75 cm in total body length and 0.5–0.75 kg body mass. They are thus the smallest known ornithischians. Fruitadens is a late-surviving member of the basal dinosaur clade Heterodontosauridae, and is the first member of this clade to be described from North America. The craniodental anatomy and diminutive body size of Fruitadens suggest that this taxon was an ecological generalist with an omnivorous diet, thus providing new insights into morphological and palaeoecological diversity within Dinosauria. Late-surviving (Late Jurassic and Early Cretaceous) heterodontosaurids are smaller and less ecologically specialized than Early (Late Triassic and Early Jurassic) heterodontosaurids, and this ecological generalization may account in part for the remarkable 100-million-year-long longevity of the clade.     

Bite traces on dicynodont bones and the early evolution of large terrestrial predators

Nied?wiedzki, G., Gorzelak, P. & Sulej, T. 2010: Bite traces on dicynodont bones and the early evolution of large terrestrial predators. Lethaia, Vol. 44, pp. 87–92. Dicynodont (Synapsida: Anomodontia) bones from the Late Triassic (late Norian/early Rhaetian) of Poland yield characteristic tooth marks that can be attributed to three ichnotaxa (Linichnus serratus, Knethichnus parallelum and Nihilichnus nihilicus). The general shape and dimension of these traces perfectly match the dental morphology of a co‐occurring carnivorous dinosaur. It is therefore concluded that early carnivorous dinosaurs were feeding on dicynodonts. This discovery constitutes one of the oldest evidence of dinosaur predator–prey interaction. It is suggested that an evolutionary increase in the size of dicynodonts across the Late Triassic may have been driven by selection pressure to reach a size refuge from early dinosaur predators. □Bite traces, dicynodonts, dinosaurs, predation, Triassic.