Cesar L. Schultz

Sequence stratigraphy of continental Triassic strata of Southernmost Brazil: a contribution to Southwestern Gondwana palaeogeography and palaeoclimate

Abstract The continental Triassic succession of southernmost Brazil comprises two second-order depositional sequences—the Sanga do Cabral (Early Triassic) and the Santa Maria (Middle to Late Triassic) supersequences. The first one includes ephemeral, low-sinuosity fluvial deposits developed on a low gradient plain. Based on fossil tetrapods, especially procolophonids, an Upper Induan age is estimated for this sequence. Facies association of the Santa Maria Supersequence indicates low-sinuosity fluvial rivers, deltas and lakes. This supersequence can be further subdivided into three third-order sequences (age provided by palaeovertebrate biostratigraphic data) as follows: Santa Maria 1 (Ladinian), Santa Maria 2 (Carnian to Early Norian) and Santa Maria 3 (probably Raethian or Early Jurassic) sequences. The Gondwanides paroxysms I and II in the Sierra de la Ventana–Cape Fold Belt are directly related to the development of both supersequences. The source area of the Sanga do Cabral Supersequence was located to the south. It consisted of an uplifted peripheral bulge situated landward of the retro-foreland system, from where older sedimentary rocks were eroded. The source area of the Santa Maria Supersequence was also positioned southwards and related to the uplifted Sul-Rio-Grandense and Uruguayan shields. The Santa Maria Supersequence stratigraphic architecture is comparable to the Triassic rift basins of Western Argentina. Diagenesis, facies and palaeontology of the studied succession suggest a dominantly semiarid climate during the Triassic.

A procolophonoid reptile with temporal fenestration from the Middle Triassic of Brazil

The small tetrapod Candelaria barbouri, from the Middle Triassic of southern Brazil, is the first example of an owenettid procolophonoid outside Africa and Madagascar. Candelaria barbouri was originally described as a primitive procolophonid; however, a re–examination of the holotype, as well as new material, reveals that C. barbouri is in fact the youngest member of the Owenettidae, extending the chronological range of the group by more than 10 million years. The recognition of C. barbouri as an owenettid points to a broader diversity and distribution for owenettids than hitherto thought. In addition, C. barbouri is the first member of the Owenettidae to exhibit temporal fenestrae, a discovery that draws attention to the significance of this feature in ‘anapsid’ reptiles.

Tapeworm Eggs in a 270 Million-Year-Old Shark Coprolite

Remains of parasites in vertebrates are rare from the Mesozoic and Paleozoic. Once most parasites that live in – or pass through – the gastrointestinal tract of vertebrates, fossil feces (coprolites) or even intestinal contents (enterolites) can eventually preserve their remains. Here we announce the discovery of a spiral shark coprolite from the Paleozoic bearing a cluster of 93 small oval-elliptical smooth-shelled structures, interpreted as eggs of a tapeworm.The eggs were found in a thin section of an elasmobranch coprolite. Most of the eggs are filled by pyrite and some have a special polar swelling (operculum), suggesting they are non-erupted eggs. One of the eggs contains a probable developing larva. The eggs are approximately 145–155 µm in length and 88–100 µm in width and vary little in size within the cluster. The depositional and morphological features of the eggs closely resemble those of cestodes. Not only do the individual eggs have features of extant tapeworms, but their deposition all together in an elongate segment is typical to modern tapeworm eggs deposited in mature segments (proglottids). This is the earliest fossil record of tapeworm parasitism of vertebrates and establishes a timeline for the evolution of cestodes. This discovery shows that the fossil record of vertebrate intestinal parasites is much older than was hitherto known and that the interaction between tapeworms and vertebrates occurred at least since the Middle-Late Permian.

A New Species Of The Late Triassic Rhynchosaur Hyperodapedon From The Santa Maria Formation Of South Brazil

A new rhynchosaur, Hyperodapedon huenei sp. nov., is described from the Upper Triassic Santa Maria Formation of the Parana Basin, Brazil. The holotype is an almost complete skull and mandible, collected at Inhamanda, in the state of Rio Grande do Sul, Brazil. The diagnosis of the genus Hyperodapedon Huxley is revised to include not only H. huxleyi Lydekker and H. gordoni Huxley (as generally accepted), but also the new species described here, various specimens usually assigned to ‘Scaphonyx fischeri’ Woodward, and ‘S’. sanjuanensis Sill. H. huenei sp. nov. exhibits a number of plesiomorphic features and appears to be the least derived species of Hyperodapedon, forming a sister taxon to the remaining members of the genus. A preliminary phylogenetic analysis for the more derived rhynchosaurs is presented. ‘Scaphonyx’sulcognathus Azevedo and Schultz represents the sister taxon of Hyperodapedon, while ‘Rhynchosaurus’spenceri Benton is considered to be a more derived Middle Triassic rhynchosaur. key words: Rhynchosauria, Hyperodapedon, Triassic, Brazil.

New stem-sauropodomorph (Dinosauria, Saurischia) from the Triassic of Brazil

Post-Triassic theropod, sauropodomorph, and ornithischian dinosaurs are readily recognized based on the set of traits that typically characterize each of these groups. On the contrary, most of the early members of those lineages lack such specializations, but share a range of generalized traits also seen in more basal dinosauromorphs. Here, we report on a new Late Triassic dinosaur from the Santa Maria Formation of Rio Grande do Sul, southern Brazil. The specimen comprises the disarticulated partial skeleton of a single individual, including most of the skull bones. Based on four phylogenetic analyses, the new dinosaur fits consistently on the sauropodomorph stem, but lacks several typical features of sauropodomorphs, showing dinosaur plesiomorphies together with some neotheropod traits. This is not an exception among basal dinosaurs, the early radiation of which is characterized by a mosaic pattern of character acquisition, resulting in the uncertain phylogenetic placement of various early members of the group.

A new specimen of Guaibasaurus candelariensis (basal Saurischia) from the Late Triassic Caturrita Formation of southern Brazil

The comparison of the anatomy of a second incomplete skeleton of Guaibasaurus candelariensis to that of basal Saurischia suggests that the origin of the “Prosauropoda” was from unknown basal saurischians, after separating from theropods (except herrerasaurids). Guaibasaurus and Saturnalia are part of this early dichotomy, and they bear mixed characters-states of basal theropods and “prosauropods” type. The Late Triassic age of both Guaibasaurus and Saturnalia (the latter is older) predates the ‘dominance’ of basal sauropodomorphs as well as the appearance of the basal theropod Zupaysaurus, both recorded from the upper Los Colorados Formation of Argentina.

Carnivorous dinocephalian from the Middle Permian of Brazil and tetrapod dispersal in Pangaea

The medial Permian (∼270–260 Ma: Guadalupian) was a time of important tetrapod faunal changes, in particular reflecting a turnover from pelycosaurian- to therapsid-grade synapsids. Until now, most knowledge on tetrapod distribution during the medial Permian has come from fossils found in the South African Karoo and the Russian Platform, whereas other areas of Pangaea are still poorly known. We present evidence for the presence of a terrestrial carnivorous vertebrate from the Middle Permian of South America based on a complete skull. Pampaphoneus biccai gen. et sp. nov. was a dinocephalian “mammal-like reptile” member of the Anteosauridae, an early therapsid predator clade known only from the Middle Permian of Russia, Kazakhstan, China, and South Africa. The genus is characterized, among other features, by postorbital bosses, short, bulbous postcanines, and strongly recurved canines. Phylogenetic analysis indicates that the Brazilian dinocephalian occupies a middle position within the Anteosauridae, reinforcing the model of a global distribution for therapsids as early as the Guadalupian. The close phylogenetic relationship of the Brazilian species to dinocephalians from South Africa and the Russian Platform suggests a closer faunistic relationship between South America and eastern Europe than previously thought, lending support to a Pangaea B-type continental reconstruction.

Digital reconstruction of the otic region and inner ear of the non-mammalian cynodont brasilitherium riograndensis (late triassic, brazil) and its relevance to the evolution of the mammalian ear

The external anatomy of the petrosal, the bony labyrinth of the inner ear, and the stapes of Brasilitherium riograndensis (specimen UFRGS-PV-1043-T) were investigated by digital 3D reconstructions based on μCT scan images. Brasilitherium is the most basal taxon bearing a distinct promontorium, although less inflated than that of Morganucodon and still lacking a flat medial facet. A bony wall formed by the petrosal separates the cochlear canal and the vestibule from the brain cavity, with an internal acoustic meatus bearing distinct foramina for the facial nerve (VII) and vestibulocochlear nerve (VIII). The semicircular canals are irregular in shape, the anterior canal being the largest and the lateral one the smallest. Brasilitherium has an elongated but straight cochlear canal. The stapes resembles the morphology of derived non-mammaliaform cynodonts, such as Probainognathus and Pachygenelus, and differs from Thrinaxodon. By the allometric relationship of the cochlear canal and the estimated body mass, Brasilitherium can be grouped with Yunnanodon and Morganucodon in a regression line, which is below the line of mammals and above the line of non-avian reptiles. Brasilitherium fits in a sequence of gradual elongation of the cochlear canal associated with the enhancement in the capacity to hear higher frequencies. Among the constraints that might have triggered these transformations in small, insectivorous, and possibly nocturnal Mesozoic cynodont taxa is the improvement of detecting acoustically active insects.

New information on the pterosaur Tupandactylus imperator, with comments on the relationships of Tapejaridae

A new specimen of Tupandactylus imperator, comprising an incomplete skull with associated lower jaw, is described. The material is the best preserved specimen of this species known so far and provides new information on the anatomy of this pterodactyloid pterosaur, especially with respect to the morphology of the lower jaw, the first one formally described for the species. Also, the new specimen shows an extensive preservation of soft tissues such as the soft-tissue component of the headcrest, ramphoteca associated with the premaxillae and lower jaw, as well as probable pycnofibres. A phylogenetic analysis was performed in order to test the relationships of the taxon within Tapejaridae. The results of the analysis support Tapejaridae, as well as monophyly of Tapejarinae and Thalassodrominae.

A reassessment of the basal dinosaur Guaibasaurus candelariensis , from the Late Triassic Caturrita Formation of south Brazil

The dinosaur record of the Santa Maria beds of Rio Grande do Sul (Mid–Late Triassic; south Brazil) includes the herrerasaur Staurikosaurus pricei , and two basal members of the sauropodomorph lineage: Saturnalia tupiniquim and Unaysaurus tolentinoi . The most enigmatic of the saurischian taxa is, however, Guaibasaurus candelariensis , previously regarded as either a basal theropod or a basal sauropodomorph. This study provides a detailed anatomical revision of all specimens originally referred to G. candelariensis by Bonaparte and co-authors, including its type-series and a more recently excavated partial postcranium. Although coming from different sites, these specimens share a unique combination of traits, and at least one possible autapomorphic feature of the pelvis, which support the inclusivity and uniqueness of the species. G. candelariensis was a medium-sized (nearly 2 m long) biped with an intriguing mix of plesiomorphic and derived (eusaurischian/theropod) features. Phylogenetic studies reveal weak support for the nesting of G. candelariensis within Theropoda, but this affinity is bolstered by various traits it shares with neotheropods. The Norian age of G. candelariensis corroborates previous studies that suggest the continuous radiation of more basal dinosauromorphs until the end of the Triassic, after the appearance of the three main dinosaur clades.