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Ricardo N. Martínez, Cecilia Apaldetti, Oscar A. Alcober, Carina E. Colombi, Paul C. Sereno, Eliana Fernandez, Paula Santi Malnis, Gustavo A. Correa, Diego Abelin
The Upper Triassic (Carnian—Norian) Ischigualasto Formation has yielded a diverse vertebrate fauna that records the initial phase of dinosaur evolution. Radioisotopic dates from ash layers within the formation provide a chrono-stratigraphic framework, and stratigraphic and sedimetological studies have subdivided the formation into four members and three abundance-based biozones. We describe two new basal dinosauromorphs, an unnamed lagerpetid and a new silesaurid, Ignotosaurus fragilis, gen. et sp. nov., which increase to 29 the number of vertebrates in the Ischigualasto fauna. We provide a census of 848 fossil specimens representing 26 vertebrate taxa logged to stratigraphic intervals of 50 m. This temporally calibrated census shows that abundance and taxonomic diversity within the Ischigualasto Formation does not change suddenly but rather appears to gradually decline in response to climatic deterioration. The only abrupt shift in faunal composition occurs at the end of the second of three biozones, when the abundant cynodont Exaeretodon is replaced by the rare dicynodont Jachaleria.
Trends of preservation are established on the basis of 385 fossil occurrences in the Ischigualasto Formation. The distribution and preservational quality of vertebrate fossils are related to sedimentary facics and stratigraphic position and follow trends in basin evolution and paleoclimate. Fossils in the Ischigualasto Formation are mainly concentrated along the margins of the basin. Floodplain deposits preserve ca. 88% of fossils, which range from unidentifiable fragments to articulated skeletons. The remaining fossil occurrences (ca. 12%) come from coarser-grained fluvial channel deposits and exhibit variable preservational quality, including some well-preserved cynodont skulls. Fossils in the basal La Peña Member (ca. 4% of occurrences) are generally highly weathered and disarticulated with hematitic permineralization. The majority of collected specimens (ca. 65% of occurrences) come from the overlying Cancha de Bochas Member and range from isolated bones to articulated skeletons. Calcite, rather than hematite, is the predominant agent of permineralization. The succeeding Valle de la Luna Member (ca. 30% of occurrences) preserves an association between vertebrate fossils and plant remains. Fossil preservation suggests prolonged exposure prior to burial under wetter conditions with superficial and penetrating precipitation of hematite. The uppermost Quebrada de la Sal Member of the Ischigualasto Formation is nearly barren of fossil remains.
Basal sauropodomorphs from the Ischigualasto Formation include Eoraptor lunensis, Panphagia protos, and Chromogisaurus novasi. Few comparisons have been made between these taxa, because Eoraptor was only recently reassessed as a basal sauropodomorph and because Panphagia and Chromogisaurus were described nearly simultaneously. We describe in detail the fully prepared bones of the holotype of Chromogisaurus novasi, examine the evidence for its taxonomic distinction, and analyze the phylogenetic relationships among basal sauropodomorphs. Our results support Chromogisaurus novasi as a valid genus and species and provide weak phylogenetic evidence favoring a series of stem taxa at the base of Sauropodomorpha. The analysis positions Panphagia as the basal-most sauropodomorph, followed by Eoraptor, Pampadromaeus, and a clade that includes Chromogisaurus and Saturnalia.
We describe a partial braincase of the basal sauropodomorph Panphagia protos from the Upper Triassic (mid-Carnian) horizons of the Ischigualasto Formation. The disarticulated braincase from a subadult individual includes one frontal, both parietals, one prootic, and the supraoccipital. The frontal is longer anteroposteriorly than it is wide transversely, has a small anterolateral process, and an elongate oval depression for the olfactory bulb. The supraoccipital is broader transversely than it is deep dorsoventrally and lacks a pronounced median nuchal eminence. Some braincase features that characterize more derived basal sauropodomorphs are not present in Panphagia, including a broader frontal and reduced anterior tympanic and floccular recesses. Panphagia appears to represent an early stage in the evolution of sauropodomorph dinosaurs.
We describe the basal sauropodomorph Eoraptor lunensis, based on the nearly complete holotypic skeleton and referred specimens, all of which were discovered in the Cancha de Bochas Member of the Ischigualasto Formation in northwestern Argentina. The lightly built skull has a slightly enlarged external naris and a spacious antorbital fossa with a prominent, everted dorsal margin and internal wall lacking any pneumatic extensions into surrounding bones. The tall quadrate is lapped along its anterior margin by the long, slender ventral process of the squamosal, and the lower jaw has a mid-mandibular joint between a tongue-shaped splenial process and a trough in the angular. All but the posterior-most maxillary and dentary crowns have a basal constriction, and the marginal denticles are larger and oriented more vertically than in typical theropod serrations. Rows of rudimentary palatal teeth are present on the pterygoid. Vertebral centra are hollow, although not demonstrably pneumatized, and all long bones have hollow shafts. The radius and ulna are more robust, the manus proportionately shorter, and the manual unguals less recurved than in the contemporaneous basal theropod Eodromaeus murphi. An outstanding feature of the manus of Eoraptor is the twisted shaft of the first phalanx of the pollex, which deflects medially the tip of the ungual as in basal sauropodomorphs. The long bones of the hind limb have more robust shafts than those of Eodromaeus, although in both genera the tibia remains slightly longer than the femur.
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