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.     

Aestivation among ornithopod dinosaurs of the African Trias

Thulborn, Richard A. 1978 07 15: Aestivation among ornithopod dinosaurs of the African Trias. Lethaia . Vol. 11, pp. 185–198. Oslo. ISSN 0024–1164.
Dental and circumstantial evidence supports the theory that ornithopod dinosaurs of the African Red Beds responded to seasonal changes in their environment by resorting to aestivation (dry season dormancy). One group of ornithopods. the heterodontosaurids. apparently suppressed tooth replacement to permit efficient grinding of their plant food and probably replaced their entire cheek dentitions while aestivating. The sympatric fabrosaurid ornithopods had a simple open-and-shut jaw action and replaced their teeth continuously in standard reptilian fashion. Seasonal dormancy must have imposed major constraints on the lives of fabrosaurids and heterodontosaurids. and these constraints are summarized in the model of a circannual life cycle.     

Anatomy and cranial functional morphology of the small-bodied dinosaur Fruitadens haagarorum from the Upper Jurassic of the USA

Background

Heterodontosaurids are an important but enigmatic and poorly understood early radiation of ornithischian dinosaurs. The late-surviving heterodontosaurid Fruitadens haagarorum from the Late Jurassic (early Tithonian) Morrison Formation of the western USA is represented by remains of several small (<1 metre total body length, <1 kg body mass) individuals that include well-preserved but incomplete cranial and postcranial material. Fruitadens is hypothesized to represent one of the smallest known ornithischian dinosaurs.

Methodology/Principal Findings

We describe the cranial and postcranial anatomy of Fruitadens in detail, providing comparisons to all other known heterodontosaurid taxa. High resolution micro-CT data provides new insights into tooth replacement and the internal anatomy of the tooth-bearing bones. Moreover, we provide a preliminary functional analysis of the skull of late-surviving heterodontosaurids, discuss the implications of Fruitadens for current understanding of heterodontosaurid monophyly, and briefly review the evolution and biogeography of heterodontosaurids.

Conclusions/Significance

The validity of Fruitadens is supported by multiple unique characters of the dentition and hindlimb as well as a distinct character combination. Fruitadens shares highly distinctive appendicular characters with other heterodontosaurids, strengthening monophyly of the clade on the basis of the postcranium. Mandibular morphology and muscle moment arms suggest that the jaws of late-surviving heterodontosaurids, including Fruitadens, were adapted for rapid biting at large gape angles, contrasting with the jaws of the stratigraphically older Heterodontosaurus, which were better suited for strong jaw adduction at small gapes. The lack of wear facets and plesiomorphic dentition suggest that Fruitadens used orthal jaw movements and employed simple puncture-crushing to process food. In combination with its small body size, these results suggest that Fruitadens was an ecological generalist, consuming select plant material and possibly insects or other invertebrates.     

The Lower Jurassic ornithischian dinosaur Heterodontosaurus tucki Crompton & Charig, 1962: cranial anatomy,functional morphology,taxonomy, and relationships

The cranial anatomy of the Lower Jurassic ornithischian dinosaur Heterodontosaurus tucki Crompton & Charig, 1962 is described in detail for the first time on the basis of two principal specimens: the holotype (SAM‐PK‐K337) and referred skull (SAM‐PK‐K1332). In addition several other specimens that have a bearing on the interpretation of the anatomy and biology of Heterodontosaurus are described. The skull and lower jaw of Heterodontosaurus are compact and robust but perhaps most notable for the heterodont dentition that merited the generic name. Details of the cranial anatomy are revealed and show that the skull is unexpectedly specialized in such an early representative of the Ornithischia, including: the closely packed, hypsodont crowns and ‘warping’ of the occlusal surfaces (created by progressive variation in the angulation of wear on successive crowns) seen in the cheek dentition; the unusual sutural relationships between the bones along the dorsal edge of the lower jaw; the very narrow, deeply vaulted palate and associated structures on the side wall of the braincase; and the indications of cranial pneumatism (more commonly seen in basal archosaurs and saurischian dinosaurs). Evidence for tooth replacement (which has long been recognized, despite frequent statements to the contrary) is suggestive of an episodic, rather than continuous, style of tooth replacement that is, yet again, unusual in diapsids generally and particularly so amongst ornithischian dinosaurs. Cranial musculature has been reconstructed and seems to conform to that typically seen in diapsids, with the exception of the encroachment of M. adductor mandibulae externus superficialis across the lateral surface of the temporal region and external surface of the lower jaw. Indications, taken from the unusual shape of the occlusal surfaces of the cheek dentition and jaw musculature, are suggestive of a novel form of jaw action in this dinosaur. The taxonomy of currently known late Karoo‐aged heterodontosaurids from southern Africa is reviewed. Although complicated by the inadequate nature of much of the known material, it is concluded that two taxa may be readily recognized: H. tucki and Abrictosaurus consors. At least one additional taxon is recognized within the taxa presently named Lanasaurus and Lycorhinus; however, both remain taxonomically problematic and their status needs to be further tested and may only be resolved by future discoveries. The only other named taxon, Geranosaurus atavus, represents an invalid name. The recognition of at least four distinct taxa indicates that the heterodontosaurids were speciose within the late Karoo ecosystem. The systematics of Heterodontosaurus and its congeners has been analysed, using a restricted sample of taxa. A basal (nongenasaurian) position within Ornithischia is re‐affirmed. There are at least four competing hypotheses concerning the phylogenetic placement of the Heterodontosauridae, so the evidence in support of the various hypotheses is reviewed in some detail. At present the best‐supported hypothesis is the one which places Heterodontosauridae in a basal (non‐genasaurian) position; however, the evidence is not fully conclusive and further information is still needed in respect of the anatomy of proximate outgroups, as well as more complete anatomical details for other heterodontosaurids. Heterodontosaurids were not such rare components of the late Karoo ecosystem as previously thought; evidence also suggests that from a phylogenetic perspective they occupied a potentially crucial position during the earliest phases of ornithischian dinosaur evolution. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011.     

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.     

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.     

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.     

The systematic position of the Triassic ornithischian dinosaur Lycorhinus angustidens

A topotype of the poorly known reptile Lycorhinus angustidens Haughton (1924) is described. This specimen demonstrates that Lycorhinus is a genuine ornithischian dinosaur of late Triassic age and not, as previously supposed, a therapsid. The generic term 'Heterodonto-saurus' Crompton & Charig (1962) is regarded as an invalid junior synonym for Lycorhinus and is discarded–though the species tucki (now referred to Lycorhinus) is retained as distinct from angustidens on the basis of slight differences in tooth structure. Lycorhinus is included within the family Hypsilophodontidae of the suborder Ornithopoda.     

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.     

A basal ceratopsian with transitional features from the Late Jurassic of northwestern China

Although the Ceratopsia and Pachycephalosauria, two major ornithischian groups, are united as the Marginocephalia, few synapomorphies have been identified due to their highly specialized body-plans. Several studies have linked the Heterodontosauridae with either the Ceratopsia or Marginocephalia, but evidence for these relationships is weak, leading most recent studies to consider the Heterodontosauridae as the basal member of another major ornithischian radiation, the Ornithopoda. Here, we report on a new basal ceratopsian dinosaur, Yinlong downsi gen. et. sp. nov. from the Late Jurassic upper part of the Shishugou Formation of Xinjiang, China. This new ceratopsian displays a series of features transitional between more derived ceratopsians and other ornithischians, shares numerous derived similarities with both the heterodontosaurids and pachycephalosaurians and provides strong evidence supporting a monophyletic Marginocephalia and its close relationship to the Heterodontosauridae. Character distributions along the marginocephalian lineage reveal that, compared to the bipedal Pachycephalosauria, which retained a primitive post-cranial body-plan, the dominantly quadrupedal ceratopsians lost many marginocephalian features and evolved their own characters early in their evolution.     

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.     

A primitive ornithischian dinosaur from the Late Triassic of South Africa, and the early evolution and diversification of Ornithischia

Although the group played an important role in the evolution of Late Mesozoic terrestrial ecosystems, the early evolutionary history of the ornithischian dinosaurs remains poorly understood. Here, we report on a new primitive ornithischian, Eocursor parvus gen. et sp. nov., from the Late Triassic (?Norian) Lower Elliot Formation of South Africa. Eocursor is known from a single specimen comprising substantial cranial and postcranial material and represents the most complete Triassic member of Ornithischia, providing the earliest evidence for the acquisition of many key ornithischian postcranial characters, including an opisthopubic pelvis. A new phylogenetic analysis positions this taxon near the base of Ornithischia, as the sister taxon to the important and diverse clade Genasauria. The problematic clade Heterodontosauridae is also positioned basal to Genasauria, suggesting that an enlarged grasping manus may represent a plesiomorphic ornithischian condition. This analysis provides additional phylogenetic support for limited ornithischian diversity during the Late Triassic, and suggests that several major ornithischian clades may have originated later than generally believed. There are few morphological differences between Late Triassic and Early Jurassic ornithischians, supporting previous suggestions that the Early Jurassic ornithischian radiation may simply represent the filling of vacant ecological space following Late Triassic terrestrial extinctions.     

The dentition of Manidens condorensis (Ornithischia; Heterodontosauridae) from the Jurassic Cañadón Asfalto Formation of Patagonia: morphology,heterodonty and the use of statistical methods for identifying isolated teeth

The recently described Manidens condorensis is one of the most completely known taxa of the family Heterodontosauridae from the southern landmasses. However, some dental aspects are not well known due to preservational problems in the type material. This contribution reports new isolated teeth found in the Cañadón Asfalto Formation (Early-Middle Jurassic). These teeth are referred to Manidens condorensis based on the presence of autapomorphic characters of the unusual dentition of this taxon, such as the highly asymmetric tooth crowns and small crenulations on each denticles. The isolated crowns are well preserved and reveal the presence of undescribed and new autapomorphical features, including apical and basal wear facets on the occlusal surface of isolated crowns and a wear surface also in the caniniform tooth. We carried out statistical analyses (including morphogeometrical and discriminant analyses), using the holotype crowns as a morphological starting point, for characterising shape variation of the crowns along the toothrow and for identifying the position of isolated crowns. These analyses allow defining morphological regions within the postcaniniform toothrow and produce a metrically based discriminant function to predict the hypothetical position of future discoveries, providing a methodological framework that could be applied to other extinct heterodont dinosaurs.     

Taxonomy,morphology, masticatory function and phylogeny of heterodontosaurid dinosaurs

Heterodontosaurids comprise an important early radiation of small-bodied herbivores that persisted for approximately 100 My from Late Triassic to Early Cretaceous time. Review of available fossils unequivocally establishes Echinodon as a very small-bodied, late-surviving northern heterodontosaurid similar to the other northern genera Fruitadens and Tianyulong. Tianyulong from northern China has unusual skeletal proportions, including a relatively large skull, short forelimb, and long manual digit II. The southern African heterodontosaurid genus Lycorhinus is established as valid, and a new taxon from the same formation is named Pegomastax africanus gen. n., sp. n. Tooth replacement and tooth-to-tooth wear is more common than previously thought among heterodontosaurids, and in Heterodontosaurus the angle of tooth-to-tooth shear is shown to increase markedly during maturation. Long-axis rotation of the lower jaw during occlusion is identified here as the most likely functional mechanism underlying marked tooth wear in mature specimens of Heterodontosaurus. Extensive tooth wear and other evidence suggests that all heterodontosaurids were predominantly or exclusively herbivores. Basal genera such as Echinodon, Fruitadens and Tianyulong with primitive, subtriangular crowns currently are known only from northern landmasses. All other genera except the enigmatic Pisanosaurus have deeper crown proportions and currently are known only from southern landmasses.     

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.     

吉林省中部早白垩世泉头组-原始鸟脚类恐龙

报道了在松辽盆地白垩纪沉积中首次发现的原始鸟脚类恐龙化石,并根据其头部特征建立一新属新种——娇小长春龙(Changchunsaurus parvus gen.etsp.nov.)。化石产于吉林省公主岭市刘房子镇山前泉头组上部紫红色含砾泥质砂岩中,同一层位还产有兽脚类、鳄类、恐龙蛋、哺乳类等化石。娇小长春龙是一种混合了原始的和衍生性状的小型鸟脚类恐龙。它具有某些比多数鸟脚类和头饰龙类要原始的特征,例如,5颗前上颌齿、前上颌骨吻部只有很短一段齿缺、前上颌骨与上颌齿之间的间隙较小、颊齿两侧的釉质对称、前上颌骨腹侧边缘与上颌骨腹侧边缘基本处于同一水平线等。同时,娇小长春龙也具有一些与真鸟脚类类似的进步特征,比如眶前孔小、外下颌孔缺失。娇小长春龙具有颧骨突,这在鸟脚类恐龙中较为罕见,它的颧骨突表面具有鲕状构造,这一特征未见于其他已知鸟脚类。娇小长春龙的前齿骨形态与角龙类接近,腹支明显长于侧支,前齿骨与齿骨的愈合方式同角龙类相似。娇小长春龙的确切系统分类位置需要进一步的工作来确定。     

Evolution of High Tooth Replacement Rates in Sauropod Dinosaurs

Background

Tooth replacement rate can be calculated in extinct animals by counting incremental lines of deposition in tooth dentin. Calculating this rate in several taxa allows for the study of the evolution of tooth replacement rate. Sauropod dinosaurs, the largest terrestrial animals that ever evolved, exhibited a diversity of tooth sizes and shapes, but little is known about their tooth replacement rates.

Methodology/Principal Findings

We present tooth replacement rate, formation time, crown volume, total dentition volume, and enamel thickness for two coexisting but distantly related and morphologically disparate sauropod dinosaurs Camarasaurus and Diplodocus. Individual tooth formation time was determined by counting daily incremental lines in dentin. Tooth replacement rate is calculated as the difference between the number of days recorded in successive replacement teeth. Each tooth family in Camarasaurus has a maximum of three replacement teeth, whereas each Diplodocus tooth family has up to five. Tooth formation times are about 1.7 times longer in Camarasaurus than in Diplodocus (315 vs. 185 days). Average tooth replacement rate in Camarasaurus is about one tooth every 62 days versus about one tooth every 35 days in Diplodocus. Despite slower tooth replacement rates in Camarasaurus, the volumetric rate of Camarasaurus tooth replacement is 10 times faster than in Diplodocus because of its substantially greater tooth volumes. A novel method to estimate replacement rate was developed and applied to several other sauropodomorphs that we were not able to thin section.

Conclusions/Significance

Differences in tooth replacement rate among sauropodomorphs likely reflect disparate feeding strategies and/or food choices, which would have facilitated the coexistence of these gigantic herbivores in one ecosystem. Early neosauropods are characterized by high tooth replacement rates (despite their large tooth size), and derived titanosaurs and diplodocoids independently evolved the highest known tooth replacement rates among archosaurs.     

Hadrosauroid Dinosaurs from the Late Cretaceous of the Sultanate of Oman

Fragmentary post-cranial remains (femora, tibia, vertebrae) of ornithischian dinosaurs from the Late Cretaceous of the Sultanate of Oman are described and referred to hadrosauroids. The specimens come from the Al-Khod Conglomerate, of latest Campanian to Maastrichtian age, in the north-eastern part of the country. Although the fragmentary condition of the fossils precludes a precise identification, various characters, including the shape of the fourth trochanter of the femur and the morphology of its distal end, support an attribution to hadrosauroids. With the possible exception of a possible phalanx from Angola, this group of ornithopod dinosaurs, which apparently originated in Laurasia, was hitherto unreported from the Afro-Arabian plate. From a paleobiogeographical point of view, the presence of hadrosauroids in Oman in all likelihood is a result of trans-Tethys dispersal from Asia or Europe, probably by way of islands in the Tethys shown on all recent paleogeographical maps of that area. Whether hadrosauroids were widespread on the Afro-Arabian landmass in the latest Cretaceous, or where restricted to the « Oman island » shown on some paleogeographical maps, remains to be determined.     

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.