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Once known solely from dental material and thought to represent an early ornithischian dinosaur, the early-diverging pseudosuchian Revueltosaurus callenderi is described from a minimum of 12 skeletons from a monodominant bonebed in the upper part of the Chinle Formation of Arizona. This material includes nearly the entire skeleton and possesses a combination of plesiomorphic and derived character states that help clarify ingroup relationships within Pseudosuchia. A phylogenetic analysis recovers R. callenderi in a clade with Aetosauria and Acaenasuchus geoffreyi that is named Aetosauriformes. Key autapomorphies of R. callenderi include a skull that is longer than the femur, a complete carapace of dermal armor including paramedian and lateral rows, as well as ventral osteoderms, and a tail end sheathed in bone. Histology of the femur and associated osteoderms demonstrate that R. callenderi was slow growing and that the individuals from the bonebed were not young juveniles but had not ceased growing. A review of other material assigned to Revueltosaurus concludes that the genus cannot be adequately diagnosed based on the type materials of the three assigned species and that only R. callenderi can be confidently referred to Revueltosaurus. Published version

Regurgitalites (fossilized regurgitates) can provide insight into the behavioral ecology and physiology of extinct species but they are rarely reported because they are difficult to identify and distinguish from coprolites. A compact mass of skeletal material from the Owl Rock Member of the Upper Triassic Chinle Formation of Arizona reveals features that identify it as a regurgitalite. Characteristics of the teeth and osteoderms in the specimen indicate that these remains belong to the pseudosuchian archosaur Revueltosaurus. Chemical and microstructural analysis revealed a dearth of gastric etching, the preservation of muscle fibers, and the absence of a phosphatic matrix, indicating that this bone mass is a regurgitalite and not a coprolite. It was probably produced by a phytosaur, rauisuchid, or temnospondyl, all of which occur in the Owl Rock Member. We offer an identification key to assist in distinguishing between different types of digestive remains produced by vertebrate carnivores.

Fossil vertebrate remains from the Keuper unit in the vicinity of the village of Krasiejów have been analyzed for almost two decades. However, the main goal of these works was focused mainly on large vertebrates. Here the authors present the first description of microvertebrate fossils from that site. The collection of around 5,000 specimens is mainly comprised of teeth and scales. The most numerous remains belong to osteichthyans: dipnoans (Ptychoceratodus and cf. Arganodus), palaeoniscids, semionotids, redfieldiids and chondrichthyans, such as Lonchidion sp., which is the first indisputable record of that genus in the Upper Triassic of Poland and the first shark at the Krasiejów locality. Tetrapod fossils consist of temnospondyl amphibians, rhynchocephalian lepidosauromorphs and archosauromorphs. Among them, temnospondyl amphibian remains are the most numerous and are represented mostly by Metoposaurus. However, on the basis of diversity in tooth morphotypes, the occurrence of other taxa cannot be excluded. Rhynchocephalians are composed of 7 fragmentary jaw morphotypes with dentition, which could indicate high taxonomic diversity (cf. Planocephalosaurus, cf. Diphydontosaurus and cf. Clevosaurus). The most varied fossil group was assigned to the archosauromorphs. The authors can distinguish at least 19 teeth morphotypes, which show similarities to the dentition of: protorosaurians (cf. Tanystropheidae), pseudosuchians (cf. Protecovasaurus, cf. Revueltosaurus), early crocodylomorphs and basal sauropodomorph dinosaurs. The first occurrence of a theropod dinosaur and cynodonts at the Krasiejów locality is also recorded. However, their remains are very rare. These new records show a high taxonomic diversity at the Krasiejów locality that contributes to our deeper understanding of Late Triassic ecosystem of Poland. publishersversion published

Acaenasuchus geoffreyi is a diminutive armored archosaur from the Upper Triassic Chinle Formation of northern Arizona, U.S.A., with uncertain evolutionary relationships and skeletal maturity. Known only from osteoderms, the taxon has been considered a valid taxon of aetosaur, juvenile specimens synonymous with the aetosaur Desmatosuchus spurensis, or a non-aetosaurian pseudosuchian archosaur. Here, we describe new fossils of Acaenasuchus geoffreyi that represent cranial, vertebral, and appendicular elements as well as previously unknown variations in the dorsal carapace and ventral shield. The skull bones are ornamented with the same anastomosing complex of ridges and grooves found on the paramedian and lateral osteoderms, and the appendicular skeleton resembles that of Revueltosaurus callenderi, Euscolosuchus olseni, aetosaurs, and other armored archosaurs such as erpetosuchids. Histology of osteoderms from the hypodigm of Acaenasuchus geoffreyi shows multiple growth lines, laminar tissue, and low vascularity, evidence that the individuals were close to skeletal maturity and not young juveniles. A revised phylogenetic analysis of early archosaurs recovers Acaenasuchus geoffreyi and Euscolosuchus olsenias sister taxa and members of a new clade that is the sister taxon of Aetosauria. This new phylogeny depicts a broader distribution of osteoderm character states previously thought to only occur in aetosaurs, demonstrating the danger of using only armor character states in aetosaur taxonomy and phylogeny. Acaenasuchus geoffreyi is also a good example of how new fossils can stabilize 'wild card' taxa in phylogenetic analyses and contributes to our understanding of the evolution of the aetosaur carapace. Petrified Forest Museum Association; Friends of Petrified Forest National Park We thank D. and J. Gillette (MNA), P. Holroyd (UCMP), and C. Levitt-Bussian (UMNH) for providing collections and archive access, and for facilitating loans from their institutions. M. Polcyn (SMU) graciously sent us locality information and photographs of SMU 75403. H.-D. Sues provided photographs of and helpful comments on Euscolosuchus olseni, and J. Strotman facilitated the molding, casting, and transfer of casts of USNM 448587 and USNM 448582. P. Holroyd (UCMP) graciously provided permission to histologically section two osteoderms, and A. Lee (Midwestern University) kindly assisted with sectioning. K. Ritterbush (University of Utah) generously provided access to her petrographic microscope imaging system, and the histological slides were imaged by N. Ong. R. Long's field notes are on file in the archives at PEFO and the UCMP and those of Charles Camp are available at the UCMP. Thanks to T. Olson for finding MDM specimens at PFV 211. We thank J. Kirkland and D. DeBlieux (Utah Geological Survey) for permission to include in this study material they collected. The collection and preparation of the specimens from PEFO was funded by the Petrified Forest Museum Association and Friends of Petrified Forest National Park. Silhouettes from phylopic.org in Figure 9 were created by S. Hartman, D. Bogdanov, and Smokeybjb and reused under the Creative Commons Attribution 3.0 Unported license (creativecommons.org/licenses/by/3.0). Thanks to V. Paes Neto and D. Gillette for constructive comments and discussion on aetosaur and glyptodont anatomy, and to A. Turner for help in constraining tree topology in TNT. Comments and suggestions made by M. B. von Baczko and I. Cerda improved the manuscript. This is Petrified Forest National Park Contribution No. 61. The conclusions presented here are those of the authors and do not represent the views of the United States Government.

As in other archosauriforms, phytosaurs and aetosaurs are characterized by the presence of well-developed osteoderms. Here we provide a comparative study on the microstructure of phytosaur (five taxa) and aetosaur (thirteen taxa) osteoderms. For outgroup comparison, we sampled osteoderms of the sister taxon to Aetosauria, Revueltosaurus callenderi, and the doswelliid Jaxtasuchus salomoni. Phytosaur, aetosaur, and Jaxtasuchus osteoderms are composed of a diploe structure, whereas the Revueltosaurus osteoderm microanatomy is more compact. The external cortex of phytosaurs, Revueltosaurus and Jaxtasuchus osteoderms is mainly composed of parallel-fibered bone. In aetosaurs, the external cortex mainly consists of lamellar bone, with lines of resorption within the primary bone indicating successive cycles of bone erosion and deposition. The basal cortex in all the specimens is composed of parallel-fibered bone, with the cancellous internal core being more strongly developed in aetosaurs than in phytosaurs. Woven or fibro-lamellar bone was recorded in both phytosaurian and aetosaurian taxa, as well as in Jaxtasuchus. Structural fibers, which at least partly suggest metaplastic origin, were only recorded in the internal core of two phytosaurs and in the basal cortex of one aetosaur. Osteoderm thickness and cancellous to compact bone ratios appear to be subject to ontogenetic change. Minimum growth mark counts in osteoderms sampled indicate that some aetosaurs and phytosaurs lived for at least two decades. Bone microstructures are more uniform in phytosaur osteoderms and show a higher level of disparity among aetosaur osteoderms, and at least in the latter, histological features are potentially apomorphic for species/genus level. Anat Rec, 297:240–260, 2014. © 2013 Wiley Periodicals, Inc.

The Moncure microvertebrate locality in the Cumnock Formation, Sanford sub-basin, North Carolina, dramatically increases the known Late Triassic age vertebrate assemblage from the Deep River Basin. The ∼50,000 recovered microvertebrate fossils include osteichthyans, amphibians, and numerous lepidosauromorph, archosauriform, and synapsid amniotes. Actinopterygian fossils consist of thousands of scales, teeth, skull, and lower jaw fragments, principally of redfieldiids and semionotids. Non-tetrapod sarcopterygians include the dipnoanArganodussp., the first record of lungfish in the Newark Supergroup. Temnospondyls are comparatively rare but the preserved centra, teeth, and skull fragments probably represent small (juvenile) metoposaurids. Two fragmentary teeth are assigned to the unusual reptileColognathus obscurus(Case). Poorly preserved but intriguing records include acrodont and pleurodont jaw fragments tentatively assigned to lepidosaurs. Among the archosauriform teeth is a taxon distinct fromR. callenderithat we assign toRevueltosaurus olseninew combination, a morphotype best assigned to cf.Galtonia, the first Newark Supergroup record ofCrosbysaurussp., and several other archosauriform tooth morphotypes, as well as grooved teeth assigned to the recently named speciesUatchitodon schneideri. Synapsids represented by molariform teeth include both “traversodontids” assigned to aff.Boreogomphodonand the “dromatheriid”Microconodon. These records are biogeographically important, with many new records for the Cumnock Formation and/or the Newark Supergroup. In particular,Colognathus,Crosbysaurus, andUatchitodonare known from basins of Adamanian age in the southwestern U.S.A. These new records include microvertebrate taxa more typical of non-Newark basins (abundant archosauriforms, temnospondyls, lungfish) as well as more typical Newark osteichthyans and synapsid-rich faunal elements.

An unusual tetrapod tooth was discovered in the Upper Triassic Chinle Formation of southeastern Utah. The tooth was originally hypothesized to pertain to Revueltosaurus, but further investigations have rejected that hypothesis. In this paper, we compare MNA V10668 to other known fossil tooth crowns from the Chinle Formation and assign the tooth to the least inclusive clade currently available, Archosauriformes, based on the presence of mesial and distal serrations, a distal keel, and a conical mesiodistal profile. Using data found in other publications and pictures of other teeth, we compare this specimen to other Triassic dental taxa. MNA V10668 shares some similarities with Crosbysaurus, Tecovasaurus, and several other named taxa, including a teardrop-shaped labiolingual profile, but possesses a unique combination of characteristics not found in other archosauromorph teeth. This increases the known diversity of archosauromorphs from the Chinle Formation and represents the first tooth of this morphotype to be found from Utah in the Late Triassic.

An unusual tetrapod tooth was discovered in the Late Triassic Chinle Formation of southeastern Utah. The tooth was originally thought to belong to Revueltosaurus but further investigations have rejected that hypothesis. In this paper we compare MNA V10668 to other known fossil teeth found in the Chinle Formation and identify the least inclusive clade it may belongs to. Using data found in other publications and pictures of other teeth, we compare this specimen to other Triassic dental taxa. MNA V10668 shares some similarities with Crosbysaurus, Tecovasaurus, and several other named taxa but possesses unique characteristics not found in other diapsid teeth. We conclude that it is most likely an archosauromorph and probably an archosauriform. This increases the known diversity of tetrapods from the Chinle Formation and represents the first tooth morphotype completely unique to Utah in the Late Triassic Period.

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.

Originally identified as an ornithischian dinosaur, Crosbysaurus harrisae has been found in New Mexico, Arizona, and its type locality in Texas, as well as in North Carolina. The genus has been reassessed by other workers in light of reinterpretations about the postcrania of another putative Triassic ornithischian, Revueltosaurus. The understanding of Triassic dental faunas has become more complicated by the extreme convergence between pseudosuchian archosaurs and ornithischian dinosaur dental morphologies. We report here on a new specimen of Crosbysaurus (MNA V10666) from the Chinle Formation at Comb Ridge in southeastern Utah. This new specimen is assigned to Crosbysaurus sp. on the basis of the unique compound posterior denticles, labiolingual width, and curvature. While MNA V10666 does not help resolve the affinities of Crosbysaurus, it does represent the extension of the geographic range of this taxon for approximately 250 kilometers. This is the first record of the genus Crosbysaurus in Utah and as such it represents the northernmost known record of this taxon. This indicates that Crosbysaurus was not limited to the southern area of the Chinle/Dockum deposition but instead was widespread across the Late Triassic paleoriver systems of western Pangea. The reported specimen was found in close association with a typical Late Triassic Chinle fauna, including phytosaurs, metoposaurs, and dinosauromorphs.

Western North America has one of the most extensive fossil records of Late Triassic dinosaurs. AllUpper Triassic strata are assigned to the Chinle Group which yields four successive, dinosaur-bearing faunas. Otischalkian (early Tuvalian) dinosaur specimens are fragmentary and indeterminate. Adamanian (late Tuvalian) dinosaurs include a herrerasaurid, Camposaurus arizonensis nov. gen. et sp., Caseosaurus crosbyensis nov. gen. et sp., Tecovasaurus murryi, a new ornithischian and an indeterminate prosauropod. Revueltian (early-middle Norian) dinosaurs include Chindesaurus bryansmalli, two new herrerasaurids, a prosauropod, Revueltosaurus callenderi, Technosaurus smalli, Lucianosaurus wildi, Protoavis texensis and the original syntypes of Coelophysis bauri. Apachean (Rhaetian) evidence for dinosaurs is principally ichnological (Grallator, Pseudotetrasauropus, Tetrasauropus), but at least three dinosaurs are known from osseous remains, including the neotype of Coelophysis bauri. Dinosaurs define three biogeographic provinces in the Late Triassic. In North America, dinosaurs are rare before the late Tuvalian (late Carnian) and appear to become increasingly numerous through the remainder of the Late Triassic.

Tooth enamel microstructure can carry significant phylogenetic, ontogenetic, and functional information within amniotes. Here we provide the first descriptions of the tooth enamel microstructure of two Late Triassic taxa, the crurotarsan Revueltosaurus callenderi Hunt and the putative ornithischian Krzyzanowskisaurus hunti (Heckert), which some consider closely related. To test the hypotheses that enamel thickness corresponds to function and/or phylogeny we analyzed the enamel of each at various scales, measuring enamel thickness and examining microstructural features throughout both longitudinal and cross-sectional thickness using previously established techniques to facilitate comparisons. Both taxa possess thick (up to ~150 µm) enamel for their size (< 20 mm crown height). Enamel in R. callenderi ranged from ~5-152 µm across a premaxillary tooth in longitudinal section, and ~42-92 µm in a maxillary/dentary tooth transverse section. K. hunti enamel thickness was ~18-155 µm longitudinally and ~29-75 µm transversely. Both also had well-developed basal unit layers (BUL) and weakly developed columnar microstructure. Well-developed lines of incremental growth (LIG) are present in both taxa, through which the columnar enamel grades into parallel crystallite enamel. Their enamel microstructure is therefore grossly similar to that of several ornithischian taxa, especially ankylosaurs, with which they are strongly convergent, and also compares well to rauisuchids and tyrannosaurids. The relatively unique combination of microstructural characteristics in the schmelzmuster of R. callenderi and K. hunti supports the hypothesis that they are closely related, but does not conclusively preclude a different taxonomic placement for K. hunti so we retain its separate generic designation.

(1990). The ornithischian form genus Revueltosaurus from the Petrified Forest of Arizona (Late Triassic: Norian; Chinle Formation) Journal of Vertebrate Paleontology: Vol. 10, 50th Anniversary - 1990, pp. 268-269.