SciCombinator

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Concept: Crurotarsi

127

Triassic predatory guild evolution reflects a period of ecological flux spurred by the catastrophic end-Permian mass extinction and terminating with the global ecological dominance of dinosaurs in the early Jurassic. In responding to this dynamic ecospace, terrestrial predator diversity attained new levels, prompting unique trophic webs with a seeming overabundance of carnivorous taxa and the evolution of entirely new predatory clades. Key among these was Crocodylomorpha, the largest living reptiles and only one of two archosaurian lineages that survive to the present day. In contrast to their existing role as top, semi-aquatic predators, the earliest crocodylomorphs were generally small-bodied, terrestrial faunivores, occupying subsidiary (meso) predator roles. Here we describe Carnufex carolinensis a new, unexpectedly large-bodied taxon with a slender and ornamented skull from the Carnian Pekin Formation (~231 Ma), representing one of the oldest and earliest diverging crocodylomorphs described to date. Carnufex bridges a problematic gap in the early evolution of pseudosuchians by spanning key transitions in bauplan evolution and body mass near the origin of Crocodylomorpha. With a skull length of >50 cm, the new taxon documents a rare instance of crocodylomorphs ascending to top-tier predator guilds in the equatorial regions of Pangea prior to the dominance of dinosaurs.

Concepts: Predation, Ecology, Extinction, Dinosaur, Triassic, Jurassic, Crocodylomorpha, Crurotarsi

33

Following the end-Permian extinction, terrestrial vertebrate diversity recovered by the Middle Triassic, and that diversity was now dominated by reptiles. However, those reptilian clades, including archosaurs and their closest relatives, are not commonly found until ~30 million years post-extinction in Late Triassic deposits despite time-calibrated phylogenetic analyses predicting an Early Triassic divergence for those clades. One of these groups from the Late Triassic, Phytosauria, is well known from a near-Pangean distribution, and this easily recognized clade bears an elongated rostrum with posteriorly retracted nares and numerous postcranial synapomorphies that are unique compared with all other contemporary reptiles. Here, we recognize the exquisitely preserved, nearly complete skeleton of Diandongosuchus fuyuanensis from the Middle Triassic of China as the oldest and basalmost phytosaur. The Middle Triassic age and lack of the characteristically-elongated rostrum fill a critical morphological and temporal gap in phytosaur evolution, indicating that the characteristic elongated rostrum of phytosaurs appeared subsequent to cranial and postcranial modifications associated with enhanced prey capture, predating that general trend of morphological evolution observed within Crocodyliformes. Additionally, Diandongosuchus supports that the clade was present across Pangea, suggesting early ecosystem exploration for Archosauriformes through nearshore environments and leading to ease of dispersal across the Tethys.

Concepts: Ecology, Bird, Cladistics, Dinosaur, Triassic, Archosaur, Crurotarsi, Phytosaur

16

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.

Concepts: Evolution, Phylogenetic nomenclature, Phylogenetics, Cladistics, Triassic, Archosaur, Crurotarsi, Aetosaur

15

A major unresolved aspect of the rise of dinosaurs is why early dinosaurs and their relatives were rare and species-poor at low paleolatitudes throughout the Late Triassic Period, a pattern persisting 30 million years after their origin and 10-15 million years after they became abundant and speciose at higher latitudes. New palynological, wildfire, organic carbon isotope, and atmospheric pCO2 data from early dinosaur-bearing strata of low paleolatitudes in western North America show that large, high-frequency, tightly correlated variations in δ(13)Corg and palynomorph ecotypes occurred within a context of elevated and increasing pCO2 and pervasive wildfires. Whereas pseudosuchian archosaur-dominated communities were able to persist in these same regions under rapidly fluctuating extreme climatic conditions until the end-Triassic, large-bodied, fast-growing tachymetabolic dinosaurian herbivores requiring greater resources were unable to adapt to unstable high CO2 environmental conditions of the Late Triassic.

Concepts: Carbon dioxide, Climate, Dinosaur, Triassic, Archosaur, Crurotarsi, Late Triassic, Thecodont

14

Crocodylomorphs originated in the Late Triassic and were the only crocodile-line archosaurs to survive the end-Triassic extinction. Recent phylogenetic analyses suggest that the closest relatives of these generally gracile, small-bodied taxa were a group of robust, large-bodied predators known as rauisuchids implying a problematic morphological gap between early crocodylomorphs and their closest relatives. Here we provide a detailed osteological description of the recently named early diverging crocodylomorph Carnufex carolinensis from the Upper Triassic Pekin Formation of North Carolina and assess its phylogenetic position within the Paracrocodylomorpha. Carnufex displays a mosaic of crocodylomorph, rauisuchid, and dinosaurian characters, as well as highly laminar cranial elements and vertebrae, ornamented dermal skull bones, a large, subtriangular antorbital fenestra, and a reduced forelimb. A phylogenetic analysis utilizing a comprehensive dataset of early paracrocodylomorphs and including seven new characters and numerous modifications to characters culled from the literature recovers Carnufex carolinensis as one of the most basal members of Crocodylomorpha, in a polytomy with two other large bodied taxa (CM 73372 and Redondavenator). The analysis also resulted in increased resolution within Crocodylomorpha and a monophyletic clade containing the holotype and two referred specimens of Hesperosuchus as well as Dromicosuchus. Carnufex occupies a key transition at the origin of Crocodylomorpha, indicating that the morphology typifying early crocodylomorphs appeared before the shift to small body size.

Concepts: Bone, Skull, Dinosaur, Triassic, Archosaur, Crocodylomorpha, Crurotarsi, Sphenosuchia

10

Archosauromorpha originated in the middle-late Permian, radiated during the Triassic, and gave rise to the crown group Archosauria, a highly successful clade of reptiles in terrestrial ecosystems over the last 250 million years. However, scientific attention has mainly focused on the diversification of archosaurs, while their stem lineage (i.e. non-archosaurian archosauromorphs) has often been overlooked in discussions of the evolutionary success of Archosauria. Here, we analyse the cranial disparity of late Permian to Early Jurassic archosauromorphs and make comparisons between non-archosaurian archosauromorphs and archosaurs (including Pseudosuchia and Ornithodira) on the basis of two-dimensional geometric morphometrics.

Concepts: Bird, Dinosaur, Triassic, Archosaur, Archosauromorpha, Crurotarsi, Rauisuchia, Avemetatarsalia

10

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.

Concepts: Arizona, Dinosaur, Triassic, Archosaur, Permian, Ornithischia, Utah, Crurotarsi

9

Dinosaurs and their close relatives grew to sizes larger than any other terrestrial animal in the history of life on Earth, and many enormous dinosaurs (e.g.,Diplodocus,Spinosaurus,Tyrannosaurus) have accessory intervertebral articulations that have been suggested to support these large body sizes. Some pseudosuchian archosaurs have been reported to have these articulations as well, but few have been characterized in these taxa because of a lower abundance of complete, three-dimensional pseudosuchian vertebral material in relation to dinosaurs. We describe the axial column of the large (∼4-5 m) poposauroid pseudosuchianPoposaurus langstonifrom the Upper Triassic of Texas (TMM Locality 31025 of the Otis Chalk localities; Dockum Group, Howard County, TX, USA).P. langstoniwas originally named from pelvic girdle elements and vertebrae; here we describe newly discovered and prepared presacral vertebrae and a presacral rib from the original excavation of the holotype in the 1940s. The well-preserved vertebrae have well-defined vertebral laminae and clear hyposphene-hypantrum intervertebral articulations, character states mentioned in pseudosuchians but rarely described. The new material demonstrates variation present in the hyposphene-hypantrum articulation through the vertebral column. We compared these morphologies to other pseudosuchians with and without the hyposphene-hypantrum articulation. Based on these careful comparisons, we provide an explicit definition for the hyposphene-hypantrum articulation applicable across Archosauria. Within Pseudosuchia, we find the hyposphene-hypantrum appeared independently in the clade at least twice, but we also see the loss of these structures in clades that had them ancestrally. Furthermore, we found the presence of large body sizes (femoral lengths >∼300 mm) and the presence of the hyposphene-hypantrum is correlated in most non-crocodylomorph pseudosuchian archosaurs with a few exceptions. This result suggests that the presence of the hyposphene-hypantrum is controlled by the increases and decreases in body size and not strictly inheritance.

Concepts: Spinal cord, Vertebral column, Bird, Human anatomy, Intervertebral disc, Dinosaur, Triassic, Crurotarsi

9

Macelognathus vagans Marsh, 1884 from the Late Jurassic Morrison Fm. of Wyoming was originally described as a dinosaur by Marsh and in 1971 Ostrom suggested crocodilian affinities. In 2005, Göhlich and collaborators identified new material of this species from Colorado as a basal crocodylomorph. However, a partial skull found in association with mandibular and postcranial remains was not described.

Concepts: Reptile, Crocodilia, Dinosaur, Archosaur, Archosauromorpha, Crocodylomorpha, Crurotarsi, Allosaurus

8

Susisuchus anatoceps is a neosuchian crocodylomorph lying outside the clade Eusuchia, and associated with the transition between basal and advanced neosuchians and the rise of early eusuchians. The specimen MPSC R1136 comprises a partially articulated postcranial skeleton and is only the third fossil assigned to this relevant taxon. Thin sections of a right rib and right ulna of this specimen have been cut for histological studies and provide the first paleohistological information of an advanced non-eusuchian neosuchian from South America. The cross-section of the ulna shows a thick cortex with 17 lines of arrested growth (LAGs), a few scattered vascular canals, and primary and secondary osteons. This bone has a free medullary cavity and a spongiosa is completely absent. Thin sections of the rib show that remodeling process was active when the animal died, with a thin cortex and a well-developed spongiosa. In the latter, few secondary osteons and 4 LAGs were identified. According to the observed data, Susisuchus anatoceps had a slow-growing histological microstructure pattern, which is common in crocodylomorphs. The high number of ulnar LAGs and the active remodeling process are indicative that this animal was at least a late subadult, at or past the age of sexual maturity. This contradicts previous studies that interpreted this and other Susisuchus anatoceps specimens as juveniles, and suggests that full-grown adults of this species were relatively small-bodied, comparable in size to modern dwarf crocodiles.

Concepts: Bone, Biology, Skeletal system, Ulna, Crocodiles, Crocodylomorpha, Crurotarsi, Eusuchia