Petrified Forest National Park

Abstract Ornithischian dinosaurs are one of the most taxonomically diverse dinosaur clades during the Mesozoic, yet their origin and early diversification remain virtually unknown. In recent years, several new Triassic ornithischian taxa have been proposed, mostly based upon isolated teeth. New discoveries of skeletal material of some of these tooth taxa indicate that these teeth can no longer be assigned to the Ornithischia using unambiguous synapomorphies. The Triassic record of ornithischian dinosaurs now comprises only three probable occurrences: Pisanosaurus and an unnamed heterodontosaurid from Argentina, and an unnamed specimen from the uppermost Triassic of South Africa. This revised Triassic record suggests that ornithischians were not very diverse or abundant through the Triassic, and there are large gaps in the Triassic ornithischian fossil record. Moreover, traditional living analogues for interpreting the feeding ecology of early ornithischians from their tooth morphology are generally inappropriate, and “herbivorous” archosaur teeth such as those found in early ornithischians are not necessarily diagnostic of herbivorous feeding.

Synopsis The North American Triassic dinosaur record has been repeatedly cited as one of the most complete early dinosaur assemblages. The discovery of Silesaurus from Poland and the recognition that Herrerasaurus and Eoraptor may not be theropods have forced a re‐evaluation of saurischian and theropod synapomorphies. Here, we re‐evaluate each purported Triassic dinosaur from North America on a specimen by specimen basis using an apomorphy‐based approach. We attempt to assign specimens to the most exclusive taxon possible. Our revision of purported Late Triassic dinosaur material from North America indicates that dinosaurs were rarer and less diverse in these strata than previously thought. This analysis concludes that non‐dinosaurian dinosauriforms were present in North America in the Late Triassic. Most of the proposed theropod specimens are fragmentary and/or indistinguishable from corresponding elements in the only well‐known Triassic theropod of North America, Coelophysis bauri. No Triassic material from North America can be assigned to Sauropodomorpha, because none of the purported ‘prosauropod’ material is diagnostic. Recent discovery of the skull and skeleton of Revueltosaurus callenderi from Arizona shows that it is a pseudosuchian archosaur, not an ornithischian dinosaur. As a result, other purported North American ornithischian teeth cannot be assigned to the Ornithischia and therefore, there are no confirmed North American Triassic ornithischians. Non‐tetanuran theropods and possible basal saurischians are the only identifiable dinosaurs recognised in North America until the beginning of the Jurassic Period.

ABSTRACT The recent discovery of early dinosauromorphs from North America demonstrates that they were contemporaries with dinosaurs and other basal archosaurs during a substantial portion of the Late Triassic Period. Hindlimb material (femora, tibiae, a fibula, astragalocalcanea, and phalanges) of Dromomeron romeri, a non-dinosauriform dinosauromorph from the Petrified Forest Member of the Chinle Formation from north-central New Mexico, is described. A new species of Dromomeron from the lower portion of the Chinle Formation (eastern Arizona) and Dockum Group (northern Texas) is also described, based on several disarticulated femora and tibiae. D. romeri, Lagerpeton, and the new taxon form the sister group to all other dinosauromorphs and demonstrate that this clade, Lagerpetidae, persisted well into the Norian. Lagerpetidae is supported by several synapomorphies: femoral head hook-shaped in medial and lateral views; ventral emargination on the anterolateral side of the femoral head; an enlarged posteromedial tuber of the proximal end of the femur; femoral crista tibiofibularis larger than the medial condyle; anteromedial corner of the distal end of the femur forms 90° or acute (>90°) angle; and a posterior ascending process of the astragalus. An ontogenetic series of the femur of Dromomeron indicates that some character states previously used in phylogenetic analyses of early dinosaurs may be ontogenetically variable.

ABSTRACT Recent stratigraphic revisions of the Upper Triassic Chinle Formation of Petrified Forest National Park, in conjunction with precise and accurate documentation of fossil tetrapod occurrences, clarified the local biostratigraphy, with regional and global implications. A significant overlap between Adamanian and Revueltian faunas is rejected, as is the validity of the Lamyan sub-land vertebrate faunachron. The Adamanian–Revueltian boundary can be precisely placed within the lower Jim Camp Wash beds of the Sonsela Member and thus does not occur at the hypothesised Tr-4 unconformity. This mid-Norian faunal turnover, may coincide with a floral turnover, based on palynology studies and also on sedimentological evidence of increasing aridity. Available age constraints bracketing the turnover horizon are consistent with the age of the Manicouagan impact event. The rise of dinosaurs in western North America did not correspond to the Adamanian–Revueltian transition, and overall dinosauromorph diversity seems to have remained at a constant level across it. The paucity of detailed Late Triassic vertebrate biostratigraphic data and radioisotopic dates makes it currently impossible to either support or reject the existence of globally synchronous Late Triassic extinctions for tetrapods.

The Newark-Hartford astrochronostratigraphic polarity timescale (APTS) was developed using a theoretically constant 405-kiloyear eccentricity cycle linked to gravitational interactions with Jupiter-Venus as a tuning target and provides a major timing calibration for about 30 million years of Late Triassic and earliest Jurassic time. While the 405-ky cycle is both unimodal and the most metronomic of the major orbital cycles thought to pace Earth's climate in numerical solutions, there has been little empirical confirmation of that behavior, especially back before the limits of orbital solutions at about 50 million years before present. Moreover, the APTS is anchored only at its younger end by U-Pb zircon dates at 201.6 million years before present and could even be missing a number of 405-ky cycles. To test the validity of the dangling APTS and orbital periodicities, we recovered a diagnostic magnetic polarity sequence in the volcaniclastic-bearing Chinle Formation in a scientific drill core from Petrified Forest National Park (Arizona) that provides an unambiguous correlation to the APTS. New high precision U-Pb detrital zircon dates from the core are indistinguishable from ages predicted by the APTS back to 215 million years before present. The agreement shows that the APTS is continuous and supports a stable 405-kiloyear cycle well beyond theoretical solutions. The validated Newark-Hartford APTS can be used as a robust framework to help differentiate provinciality from global temporal patterns in the ecological rise of early dinosaurs in the Late Triassic, amongst other problems.

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.

Abstract Aetosauria is a clade of obligately quadrupedal, heavily armoured pseudosuchians known from Upper Triassic (late Carnian–Rhaetian) strata on every modern continent except Australia and Antarctica. As many as 22 genera and 26 species ranging from 1 to 6 m in length, and with a body mass ranging from less than 10 to more than 500 kg, are known. Aetosauroides scagliai was recently recovered as the most basal aetosaur, placed outside of Stagonolepididae (the last common ancestor of Desmatosuchus and Aetosaurus ). Interrelationships among the basal aetosaurs are not well understood but two clades with relatively apomorphic armour – the spinose Desmatosuchinae and the generally wide-bodied Typothoracisinae – are consistently recognized. Paramedian and lateral osteoderms are often distinctive at the generic level but variation within the carapace is not well understood in many taxa, warranting caution in assigning isolated osteoderms to specific taxa. The aetosaur skull and dentition varies across taxa, and there is increasing evidence that at least some aetosaurs relied on invertebrates and/or small vertebrates as a food source. Histological evidence indicates that, after an initial period of rapid growth, lines of arrested growth (LAGs) are common and later growth was relatively slow. The common and widespread Late Triassic ichnogenus Brachychirotherium probably represents the track of an aetosaur.

Synopsis Study of aetosaurian archosaur material demonstrates that the dermal armour of Desmatosuchus chamaensis shares almost no characters with that of Desmatosuchus haplocerus. Instead, the ornamentation and overall morphology of the lateral and paramedian armour of ’D.’ chamaensis most closely resembles that of typothoracisine aetosaurs such as Paratypothorax. Autapomorphies of ’D.’ chamaensis, for example the extension of the dorsal eminences of the paramedian plates into elongate, recurved spikes, warrant generic distinction for this taxon. This placement is also supported by a new phylogenetic hypothesis for the Aetosauria in which ’D.’ chamaensis is a sister taxon of Paratypothorax and distinct from Desmatosuchus. Therefore, a new genus, Heliocanthus is erected for ’D.’ chamaensis. Past phylogenetic hypotheses of the Aetosauria have been plagued by poorly supported topologies, coding errors and poor character construction. A new hypothesis places emphasis on characters of the lateral dermal armour, a character set previously under‐utilised. Detailed examination of aetosaur material suggests that the aetosaurs can be divided into three groups based on the morphology of the lateral armour. Whereas it appears that the characters relating to the ornamentation of the paramedian armour are homoplastic, those relating to the overall morphology of the lateral armour may possess a stronger phylogenetic signal.

Abstract Dilophosaurus wetherilli was the largest animal known to have lived on land in North America during the Early Jurassic. Despite its charismatic presence in pop culture and dinosaurian phylogenetic analyses, major aspects of the skeletal anatomy, taxonomy, ontogeny, and evolutionary relationships of this dinosaur remain unknown. Skeletons of this species were collected from the middle and lower part of the Kayenta Formation in the Navajo Nation in northern Arizona. Redescription of the holotype, referred, and previously undescribed specimens of Dilophosaurus wetherilli supports the existence of a single species of crested, large-bodied theropod in the Kayenta Formation. The parasagittal nasolacrimal crests are uniquely constructed by a small ridge on the nasal process of the premaxilla, dorsoventrally expanded nasal, and tall lacrimal that includes a posterior process behind the eye. The cervical vertebrae exhibit serial variation within the posterior centrodiapophyseal lamina, which bifurcates and reunites down the neck. Iterative specimen-based phylogenetic analyses result in each of the additional specimens recovered as the sister taxon to the holotype. When all five specimens are included in an analysis, they form a monophyletic clade that supports the monotypy of the genus. Dilophosaurus wetherilli is not recovered as a ceratosaur or coelophysoid, but is instead a non-averostran neotheropod in a grade with other stem-averostrans such as Cryolophosaurus ellioti and Zupaysaurus rougieri . We did not recover a monophyletic ‘Dilophosauridae.’ Instead of being apomorphic for a small clade of early theropods, it is more likely that elaboration of the nasals and lacrimals of stem-averostrans is plesiomorphically present in early ceratosaurs and tetanurans that share those features. Many characters of the axial skeleton of Dilophosaurus wetherilli are derived compared to Late Triassic theropods and may be associated with macropredation and an increase in body size in Theropoda across the Triassic-Jurassic boundary.

BACKGROUND: Recent revisions to the Sonsela Member of the Chinle Formation in Petrified Forest National Park have presented a three-part lithostratigraphic model based on unconventional correlations of sandstone beds. As a vertebrate faunal transition is recorded within this stratigraphic interval, these correlations, and the purported existence of a depositional hiatus (the Tr-4 unconformity) at about the same level, must be carefully re-examined. METHODOLOGY/PRINCIPAL FINDINGS: Our investigations demonstrate the neglected necessity of walking out contacts and mapping when constructing lithostratigraphic models, and providing UTM coordinates and labeled photographs for all measured sections. We correct correlation errors within the Sonsela Member, demonstrate that there are multiple Flattops One sandstones, all of which are higher than the traditional Sonsela sandstone bed, that the Sonsela sandstone bed and Rainbow Forest Bed are equivalent, that the Rainbow Forest Bed is higher than the sandstones at the base of Blue Mesa and Agate Mesa, that strata formerly assigned to the Jim Camp Wash beds occur at two stratigraphic levels, and that there are multiple persistent silcrete horizons within the Sonsela Member. CONCLUSIONS/SIGNIFICANCE: We present a revised five-part model for the Sonsela Member. The units from lowest to highest are: the Camp Butte beds, Lot's Wife beds, Jasper Forest bed (the Sonsela sandstone)/Rainbow Forest Bed, Jim Camp Wash beds, and Martha's Butte beds (including the Flattops One sandstones). Although there are numerous degradational/aggradational cycles within the Chinle Formation, a single unconformable horizon within or at the base of the Sonsela Member that can be traced across the entire western United States (the "Tr-4 unconformity") probably does not exist. The shift from relatively humid and poorly-drained to arid and well-drained climatic conditions began during deposition of the Sonsela Member (low in the Jim Camp Wash beds), well after the Carnian-Norian transition.

ABSTRACT The Post Quarry, within the lower part of the type section of the Upper Triassic Cooper Canyon Formation in southern Garza County, western Texas, contains a remarkably diverse vertebrate assemblage. The Post Quarry has produced: the small temnospondyl Rileymillerus cosgriffi ; the metoposaurid Apachesaurus gregorii ; possible dicynodonts and eucynodonts; a clevosaurid sphenodontian; non-archosauriform archosauromorphs ( Trilophosaurus dornorum , simiosaurians, and possibly Malerisaurus ); the phytosaur Leptosuchus ; several aetosaurs ( Calyptosuchus wellesi , Typothorax coccinarum , Paratypothorax , and Desmatosuchus smalli ); the poposauroid Shuvosaurus inexpectatus (“ Chatterjeea elegans ”); the rauisuchid Postosuchus kirkpatricki ; an early crocodylomorph; several dinosauromorphs (the lagerpetid Dromomeron gregorii , the silesaurid Technosaurus smalli , a herrerasaurid, and an early neotheropod); and several enigmatic small diapsids. Revised lithostratigraphic correlations of the lower Cooper Canyon Formation with the Tecovas Formation, the occurrence of Leptosuchus , and the overall composition of the assemblage indicate that the Post Quarry falls within the Adamanian biozone, and not the Revueltian biozone. Stratigraphic subdivision of the Adamanian biozone may be possible, and the Post Quarry may be correlative with the upper part of the Adamanian biozone in Arizona. The age of the Post Quarry assemblage is possibly late Lacian or earliest Alaunian (late early Norian or earliest middle Norian), between 220 and 215 Ma.

Aetosauria is an early-diverging clade of pseudosuchians (crocodile-line archosaurs) that had a global distribution and high species diversity as a key component of various Late Triassic terrestrial faunas. It is one of only two Late Triassic clades of large herbivorous archosaurs, and thus served a critical ecological role. Nonetheless, aetosaur phylogenetic relationships are still poorly understood, owing to an overreliance on osteoderm characters, which are often poorly constructed and suspected to be highly homoplastic. A new phylogenetic analysis of the Aetosauria, comprising 27 taxa and 83 characters, includes more than 40 new characters that focus on better sampling the cranial and endoskeletal regions, and represents the most comprenhensive phylogeny of the clade to date. Parsimony analysis recovered three most parsimonious trees; the strict consensus of these trees finds an Aetosauria that is divided into two main clades: Desmatosuchia, which includes the Desmatosuchinae and the Stagonolepidinae, and Aetosaurinae, which includes the Typothoracinae. As defined Desmatosuchinae now contains Neoaetosauroides engaeus and several taxa that were previously referred to the genus Stagonolepis, and a new clade, Desmatosuchini, is erected for taxa more closely related to Desmatosuchus. Overall support for some clades is still weak, and Partitioned Bremer Support (PBS) is applied for the first time to a strictly morphological dataset demonstrating that this weak support is in part because of conflict in the phylogenetic signals of cranial versus postcranial characters. PBS helps identify homoplasy among characters from various body regions, presumably the result of convergent evolution within discrete anatomical modules. It is likely that at least some of this character conflict results from different body regions evolving at different rates, which may have been under different selective pressures.

Abstract The Upper Triassic Chinle Formation is a critical non-marine archive of low-paleolatitude biotic and environmental change in southwestern North America. The well-studied and highly fossiliferous Chinle strata at Petrified Forest National Park (PFNP), Arizona, preserve a biotic turnover event recorded by vertebrate and palynomorph fossils, which has been alternatively hypothesized to coincide with tectonically driven climate change or with the Manicouagan impact event at ca. 215.5 Ma. Previous outcrop-based geochronologic age constraints are difficult to put in an accurate stratigraphic framework because lateral facies changes and discontinuous outcrops allow for multiple interpretations. A major goal of the Colorado Plateau Coring Project (CPCP) was to retrieve a continuous record in unambiguous superposition designed to remedy this situation. We sampled the 520-m-long core 1A of the CPCP to develop an accurate age model in unquestionable superposition by combining U-Pb zircon ages and magnetostratigraphy. From 13 horizons of volcanic detritus-rich siltstone and sandstone, we screened up to ∼300 zircon crystals per sample using laser ablation–inductively coupled plasma–mass spectrometry and subsequently analyzed up to 19 crystals of the youngest age population using the chemical abrasion–isotope dilution–thermal ionization mass (CA-ID-TIMS) spectrometry method. These data provide new maximum depositional ages for the top of the Moenkopi Formation (ca. 241 Ma), the lower Blue Mesa Member (ca. 222 Ma), and the lower (ca. 218 to 217 Ma) and upper (ca. 213.5 Ma) Sonsela Member. The maximum depositional ages obtained for the upper Chinle Formation fall well within previously proposed age constraints, whereas the maximum depositional ages for the lower Chinle Formation are relatively younger than previously proposed ages from outcrop; however, core to outcrop stratigraphic correlations remain uncertain. By correlating our new ages with the magnetostratigraphy of the core, two feasible age model solutions can be proposed. Model 1 assumes that the youngest, coherent U-Pb age clusters of each sample are representative of the maximum depositional ages and are close to (<1 Ma difference) the true time of deposition throughout the Sonsela Member. This model suggests a significant decrease in average sediment accumulation rate in the mid-Sonsela Member. Hence, the biotic turnover preserved in the mid-Sonsela Member at PFNP is also middle Norian in age, but may, at least partially, be an artifact of a condensed section. Model 2 following the magnetostratigraphic-based age model for the CPCP core 1A suggests instead that the ages from the lower and middle Sonsela Member are inherited populations of zircon crystals that are 1–3 Ma older than the true depositional age of the strata. This results in a model in which no sudden decrease in sediment accumulation rate is necessary and implies that the base of the Sonsela Member is no older than ca. 216 Ma. Independent of these alternatives, both age models agree that none of the preserved Chinle Formation in PFNP is Carnian (>227 Ma) in age, and hence the biotic turnover event cannot be correlated to the Carnian–Norian boundary but is rather a mid-Norian event. Our age models demonstrate the powers, but also the challenges, of integrating detrital CA-ID-TIMS ages with magnetostratigraphic data to properly interpret complex sedimentary sequences.

ABSTRACT A small aetosaur skeleton collected in 1939 from the Tecovas Formation of Texas and assigned to Desmatosuchus is reassigned to a new taxon, Sierritasuchus macalpini. Phylogenetic analysis suggests that Sierritasuchus is a member of the Desmatosuchinae. It can be distinguished from other desmatosuchines by two autapomorphies: (1) recurved spines on the lateral plates that are triangular in cross-section with a sharply ridged anterior edge; and (2) the presence of a sharp, ventrally oriented ridge on the posterior faces of the dorsal eminences of the paramedian plates, as well as a unique combination of characters including the presence of an anterior bar on the paramedian and lateral plates, a random pattern of ornamentation on the paramedian plates, and a dorsal eminence that contacts the posterior plate margin of the paramedian plates. Histological study of the holotypic plates in combination with comparison to a growth series in Typothorax, and size-independent growth indicators such as neurocentral suture closure suggests that the specimen is neither a young juvenile nor a fully-grown adult. The recognition of morphologically distinct specimens such as the holotype and referred material of Sierritasuchus demonstrates that past practices of assigning aetosaur specimens to known taxa based on superficial resemblance has masked diversity in this clade. Voucher specimens for biochronologic and biogeographic analyses should be carefully investigated before being used for such studies.

Crown-group frogs (Anura) originated over 200 Ma according to molecular phylogenetic analyses, though only a few fossils from high latitudes chronicle the first approximately 60 Myr of frog evolution and distribution. We report fossils that represent both the first Late Triassic and the earliest equatorial record of Salientia, the group that includes stem and crown-frogs. These small fossils consist of complete and partial ilia with anteriorly directed, elongate and distally hollow iliac blades. These features of these ilia, including the lack of a prominent dorsal protuberance and a shaft that is much longer than the acetabular region, suggest a closer affinity to crown-group Anura than to Early Triassic stem anurans Triadobatrachus from Madagascar and Czatkobatrachus from Poland, both high-latitude records. The new fossils demonstrate that crown anurans may have been present in the Late Triassic equatorial region of Pangea. Furthermore, the presence of Early Jurassic anurans in the same stratigraphic sequence ( Prosalirus bitis from the Kayenta Formation) suggests that anurans survived the climatic aridification of this region in the early Mesozoic. These fossils highlight the importance of the targeted collection of microfossils and provide further evidence for the presence of crown-group representatives of terrestrial vertebrates prior to the end-Triassic extinction.

Sarahsaurus aurifontanalis, from the Kayenta Formation of Arizona, is one of only three sauropodomorph dinosaurs known from the Early Jurassic of North America. It joins Anchisaurus polyzelus, from the older Portland Formation of the Hartford Basin, and Seitaad reussi, from the younger Navajo Sandstone of Utah, in representing the oldest North American sauropodomorphs. If it is true that sauropodomorphs were absent from North America during the Late Triassic, the relationship among these three dinosaurs offers a test of the mechanisms that drove recovery in North American biodiversity following the end-Triassic extinction event. Here we provide the first thorough description of Sarahsaurus aurifontanalis based on completed preparation and computed tomographic imaging of the holotype and referred specimens. With new anatomical data, our phylogenetic analysis supports the conclusion that Sarahsaurus aurifontanalis is nested within the primarily Gondwanan clade Massospondylidae, while agreeing with previous analyses that the three North American sauropodomorphs do not themselves form an exclusive clade. A revised diagnosis and more thorough understanding of the anatomy of Sarahsaurus aurifontanalis support the view that independent dispersal events were at least partly responsible for the recovery in North American vertebrate diversity following a major extinction event.

The Upper Triassic Chinle Formation in southwestern Laurentia is the oldest distinctive record of Early Mesozoic Cordilleran arc magmatism, in the form of detrital zircons and volcanic clasts. Initial deposition of the basal Shinarump and Mesa Redondo members, herein collectively called the Shinarump conglomerate, began in Late Triassic time, yet the earliest known arc magmatism is older by as much as 40 m.y.

Abstract Living amphibians (Lissamphibia) include frogs and salamanders (Batrachia) and the limbless worm-like caecilians (Gymnophiona). The estimated Palaeozoic era gymnophionan–batrachian molecular divergence 1 suggests a major gap in the record of crown lissamphibians prior to their earliest fossil occurrences in the Triassic period 2–6 . Recent studies find a monophyletic Batrachia within dissorophoid temnospondyls 7–10 , but the absence of pre-Jurassic period caecilian fossils 11,12 has made their relationships to batrachians and affinities to Palaeozoic tetrapods controversial 1,8,13,14 . Here we report the geologically oldest stem caecilian—a crown lissamphibian from the Late Triassic epoch of Arizona, USA—extending the caecilian record by around 35 million years. These fossils illuminate the tempo and mode of early caecilian morphological and functional evolution, demonstrating a delayed acquisition of musculoskeletal features associated with fossoriality in living caecilians, including the dual jaw closure mechanism 15,16 , reduced orbits 17 and the tentacular organ 18 . The provenance of these fossils suggests a Pangaean equatorial origin for caecilians, implying that living caecilian biogeography reflects conserved aspects of caecilian function and physiology 19 , in combination with vicariance patterns driven by plate tectonics 20 . These fossils reveal a combination of features that is unique to caecilians alongside features that are shared with batrachian and dissorophoid temnospondyls, providing new and compelling evidence supporting a single origin of living amphibians within dissorophoid temnospondyls.

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.

Abstract. Uranium–lead (U–Pb) geochronology was conducted by laser ablation – inductively coupled plasma mass spectrometry (LA-ICPMS) on 7175 detrital zircon grains from 29 samples from the Coconino Sandstone, Moenkopi Formation, and Chinle Formation. These samples were recovered from ∼ 520 m of drill core that was acquired during the Colorado Plateau Coring Project (CPCP), located in Petrified Forest National Park (Arizona). A sample from the lower Permian Coconino Sandstone yields a broad distribution of Proterozoic and Paleozoic ages that are consistent with derivation from the Appalachian and Ouachita orogens, with little input from local basement or Ancestral Rocky Mountain sources. Four samples from the Holbrook Member of the Moenkopi Formation yield a different set of Precambrian and Paleozoic age groups, indicating derivation from the Ouachita orogen, the East Mexico arc, and the Permo-Triassic arc built along the Cordilleran margin. A total of 23 samples from the Chinle Formation contain variable proportions of Proterozoic and Paleozoic zircon grains but are dominated by Late Triassic grains. LA-ICPMS ages of these grains belong to five main groups that correspond to the Mesa Redondo Member, Blue Mesa Member and lower part of the Sonsela Member, upper part of the Sonsela Member, middle part of the Petrified Forest Member, and upper part of the Petrified Forest Member. The ages of pre-Triassic grains also correspond to these chronostratigraphic units and are interpreted to reflect varying contributions from the Appalachian orogen to the east, Ouachita orogen to the southeast, Precambrian basement exposed in the ancestral Mogollon Highlands to the south, East Mexico arc, and Permian–Triassic arc built along the southern Cordilleran margin. Triassic grains in each chronostratigraphic unit also have distinct U and thorium (Th) concentrations, which are interpreted to reflect temporal changes in the chemistry of arc magmatism. Comparison of our LA-ICPMS ages with available chemical abrasion thermal ionization mass spectrometry (CA-TIMS) ages and new magnetostratigraphic data provides new insights into the depositional history of the Chinle Formation, as well as methods utilized to determine depositional ages of fluvial strata. For parts of the Chinle Formation that are dominated by fine-grained clastic strata (e.g., mudstone and siltstone), such as the Blue Mesa Member and Petrified Forest Member, all three chronometers agree (to within ∼ 1 Myr), and robust depositional chronologies have been determined. In contrast, for stratigraphic intervals dominated by coarse-grained clastic strata (e.g., sandstone), such as most of the Sonsela Member, the three chronologic records disagree due to recycling of older zircon grains and variable dilution of syn-depositional-age grains. This results in LA-ICPMS ages that significantly predate deposition and CA-TIMS ages that range between the other two chronometers. These complications challenge attempts to establish a well-defined chronostratigraphic age model for the Chinle Formation.