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Aetosauria: A clade of heavily armored Upper Triassic archosaurs

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Andrew B. Heckert
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Aetosaur fossils from the Upper Triassic Ischigualasto Formation of Argentina formerly assigned to Aetosauroides scagliai Casamiquela and Argentinosuchus bonapartei Casamiquela are reassigned to Stagonolepis robertsoni Agassiz (small specimens) and S. wellesi (Long and Ballew) (larger specimens). Numerous features of the skull, vertebral column, appendicular skeleton, and particularly the armor, of these Argentinian aetosaurs are identical to those of Stagonolepis and differ from other aetosaurs. Identification of Stagonolepis in South America has important implications for the correlation of nonmarine strata across Late Triassic Pangea. Stagonolepis is an index taxon of the Adamanian land-vertebrate faunachron of latest Carnian age. Its occurrences in the Ischigualasto and Santa Maria formations, as well as at the type locality in the Lossiemouth Sandstone of Scotland and in the Blasensandstein of the German Keuper, cross-correlate with a well-established tetrapod biostratigraphy of the Chinle Group in western North America. Thus, all Stagonolepis records are of Adamanian (latest Carnian) age, not Otischalkian (early or “middle” Carnian) age, as proposed by some earlier workers. This correlation also demonstrates that the oldest known dinosaurs are not from the Ischigualasto and Santa Maria formations. Previous workers obtained radioisotopic dates of 227.8 Ma from the Ischigualasto Formation, providing a maximum numerical date for Stagonolepis -bearing units. This date must be late Carnian, not of Ladinian age, and thus supports the apparent age of the Ladinian-Carnian boundary (232 Ma) of time scales based on the Newark Supergroup in eastern North America.
Andrew B. Heckert
added 5 research items
We describe a new species of the aetosaur Coahomasuchus , C . chathamensis , based on an incomplete, but largely articulated, anterior portion of a skeleton recovered from a quarry in the Upper Triassic Pekin Formation of Chatham County, North Carolina. This is only the second documented occurrence of Coahomasuchus , with the other being the holotype of C . kahleorum Heckert and Lucas, 1999 from the Upper Triassic Colorado City Formation of Texas. Although much of the specimen is the same size as the holotype of C . kahleorum , the dorsal paramedian osteoderms of the North Carolina taxon are considerably (~1.3×) wider than homologous counterparts in C . kahleorum , and the ventral thoracic osteoderms are also rectangular (~1.5× wider than long), rather than square, presumably to accommodate the wider body. This is a rare instance where two articulated specimens of closely related aetosaur species are available for direct comparison of homologous osteoderms. Isolated osteoderms with similar ornamentation from the same locality indicate that C . chathamensis may have been one of the earliest aetosaurs to attain the broad osteoderm proportions (width:length >3.5:1) otherwise known solely from later branching, spinose taxa such as Typothorax . The co-occurrence of Lucasuchus and Coahomasuchus in both North Carolina and Texas supports past correlations indicating an Otischalkian (Carnian) age for these strata and demonstrates that plesiomorphic, non-spinose aetosaur genera were not necessarily endemic to a single basin in North America.
Aetosauria is a clade of heavily armored, quadrupedal omnivorous to herbivorous archosaurs known from the Late Triassic across what was the supercontinent of Pangea. Their abundance in many deposits relative to the paucity of other Triassic herbivores indicates that they were key components of Late Triassic ecosystems. However, their evolutionary relationships remain contentious due, in large part, to their extensive dermal armor, which often obstructs observation of internal skeletal anatomy and limits access to potentially informative characters. In an attempt to address this problem we reanalyzed the holotype of a recently described species of Coahomasuchus , C. chathamensis , from the Sanford sub-basin of North Carolina using computed tomography (CT). CT scans of the holotype specimen clarify preservation of the skeleton, revealing several articulated vertebrae and ribs, an isolated vertebra, left ulna, left scapula, and the right humerus, though none of the material resulted in updated phylogenetic scorings. Reexamination of aetosaur materials from the holotype locality also indicates that several isolated osteoderms and elements of the appendicular skeleton are newly referable. Based on these results, we update the Coahomasuchus chathamensis hypodigm and conduct a revised phylogenetic analysis with improved character scorings for Coahomasuchus and several other aetosaurs. Our study recovers Coahomasuchus in a polytomy with Aetosaurus and the Typothoracinae, in contrast with a recent analysis that recovered Coahomasuchus as a wild-card taxon.
Data matrix used for TNT analysis Complete set of character scorings for taxa used for the phylogenetic analysis in this study. Provided in .txt format for ease of editing and use in multiple programs.