William G. Parker

Early ornithischian dinosaurs: the Triassic record

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.

A critical re‐evaluation of the Late Triassic dinosaur taxa of North America

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.

Hindlimb Osteology and Distribution of Basal Dinosauromorphs from the Late Triassic of North America

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.

Reassessment of the Aetosaur ‘Desmatosuchus’ chamaensis with a reanalysis of the phylogeny of the Aetosauria (Archosauria: Pseudosuchia)

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.

The Late Triassic (Norian) Adamanian–Revueltian tetrapod faunal transition in the Chinle Formation of Petrified Forest National Park, Arizona

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.

Revised Lithostratigraphy of the Sonsela Member (Chinle Formation, Upper Triassic) in the Southern Part of Petrified Forest National Park, Arizona

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, Lots Wife beds, Jasper Forest bed (the Sonsela sandstone)/Rainbow Forest Bed, Jim Camp Wash beds, and Marthas 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.

Aetosauria: a clade of armoured pseudosuchians from the Upper Triassic continental beds

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.

A New Desmatosuchine Aetosaur (Archosauria: Suchia) from the Upper Triassic Tecovas Formation (Dockum Group) of Texas

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.

Using positional homology in aetosaur (Archosauria: Pseudosuchia) osteoderms to evaluate the taxonomic status of Lucasuchus hunti

ABSTRACT The Otis Chalk quarries in the Upper Triassic Dockum Group of West Texas have produced aetosaur material that most workers have suggested represents two distinct morphotypes. We use characters from aetosaur specimens with articulated or semi-articulated carapaces in which the anteroposterior placement of osteoderms can be established with certainty to compare homologous osteoderms in the Otis Chalk material. This study confirms that the genera Longosuchus and Lucasuchus are distinct morphotypes, which differ in that the former taxon has paramedian osteoderms with random pitted ornamentation and low pyramidal bosses that contact the posterior margin, and spines on the lateral osteoderms that are posteriorly emarginated, whereas the latter taxon has paramedians with a strongly radial ornamentation and large conical eminences, and spines on the lateral osteoderms that are not posteriorly emarginated. Both taxa also have paramedians that are overlapped anteriorly by the laterals, a character that may be a synapomorphy of desmatosuchine aetosaurs. The arguments that these morphotypes represent ontogenetic stages or sexual dimorphs of a single biological species cannot be corroborated using either comparisons with modern pseudosuchians, other aetosaur taxa, or stratigraphic ranges. Longosuchus is known only from the type area and has no utility as an index taxon of the Otischalkian land-vertebrate faunachron, although Lucasuchus suggests a tentative correlation between part of the Dockum Group of Texas and the Pekin Formation of North Carolina.

Revised phylogenetic analysis of the Aetosauria (Archosauria: Pseudosuchia); assessing the effects of incongruent morphological character sets

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.