Early Norian Research Articles

The biogeographic history of the allokotosaurian archosauromorphs in the Triassic of Pangaea

ABSTRACT Allokotosauria (Trilophosauridae + Azendohsauridae) is a clade of Early – Late Triassic quadruped, sprawling archosauromorphs with diverse dietary habits, known from Madagascar, India, Morocco, North America, and Europe. We conducted the first quantitative biogeographic analysis of the clade to reconstruct its ancestral areas and dispersal events. The results estimated the India + Tanzania area as ancestral for Allokotosauria. Eastern North America was reconstructed as the ancestral area of Trilophosauridae, implying an Early Triassic Southern to Northern Hemisphere dispersal. Subsequently, trilophosaurids were restricted to the Northern Hemisphere, and at least two dispersal events from Europe to western North America are estimated during the early Anisian and probably during the early Carnian, respectively. Azendohsauridae retained the ancestral area of India + Tanzania during much of its evolutionary history. Two dispersal events to western and eastern North America, respectively, are reconstructed during the Carnian – early Norian, likely during or in the aftermath of the Carnian Pluvial Episode (CPE). This pattern mirrors that of early dinosaurs, which were restricted to the Southern Hemisphere with dispersals to the Northern Hemisphere not recorded until the end of the CPE. This study informs about biogeographic patterns involved in the biotic recovery after the end-Permian mass extinction and the early evolutionary radiation of Archosauromorpha.

Biostratigraphic significance of a new record of Protuberum cabralense, a bizarre traversodontid cynodont from the Middle‑Late Triassic of Southern Brazil

ABSTRACT Traversodontid cynodonts are one of the most abundant and diverse groups of tetrapods in the Triassic assemblages of Brazil. These cynodonts are distributed across four Brazilian Assemblage Zones (AZ), ranging from the late Ladinian to the early Norian. The most taxonomically diverse record comes from the Dinodontosaurus AZ, which has so far yielded at least six traversodontid genera. Among these taxa, Protuberum cabralense represents one of the most bizarre forms. This cynodont is characterised by the presence of enlarged and rounded protuberances along the dorsal surface of the ribs, the vertebral neural spines and dorsal portion of the iliac blades. These structures have been suggested to serve as a defensive mechanism or related to burrowing/digging habits. We report the first occurrence of P. cabralense at the Linha Várzea 1 site, located in the municipality of Paraíso do Sul, Brazil. This site is considered late Ladinian to early Carnian in age. The new material consists of five rib fragments, which possess the characteristic protuberances on their dorsal surface. A biostratigraphic investigation demonstrates the absence of P. cabralense in fossiliferous sites that yielded the cynodonts Luangwa. This observation contributes to the discussion regarding a potential subdivision within the Dinodontosaurus AZ.

Sedimentary facies and mineral provenance of Upper Triassic sandstones offshore Kvitøya, Svalbard: implications for palaeogeographic interpretations in the northern Barents Shelf area

Upper Triassic (Carnian) sandstones of the De Geerdalen Formation cored south of the island of Kvitøya (80°N), north-easternmost Svalbard, are described in terms of sedimentary facies and petrography and compared regionally in the northern Barents Shelf. The succession off Kvitøya is characterized by its great thickness and is dominated by deltaic deposits with high sand content of lithic–feldspathic compositions. Comparison of sediment facies and sandstone compositions with adjacent areas suggest that the succession off Kvitøya is part of a larger delta system with its main sediment source from the east. The delta sedimentation was terminated by marine transgression in the earliest Norian. The sandstone compositions off Kvitøya differ from nearby locations by the higher content of cherty rock fragments and reworked volcanic debris in the Kvitøya sandstone, which is most distinct in the lower part of the succession. Provenance signatures are investigated by mineral–chemical analysis of detrital feldspars, rock fragments, garnet and Cr-spinel, characterizing a wide variety of igneous, metamorphic and sedimentary terranes, including palaeo-Urals and areas farther to east. Additional, more proximal sediment source areas may also have existed that could explain the increased sediment thickness and the mineralogical immature sandstone compositions of the Carnian sediments off Kvitøya.

Osteology of the sauropodomorph dinosaur Jaklapallisaurus asymmetricus from the Late Triassic of central India.

The Gondwana formations exposed in the Pranhita-Godavari Valley of central India include Middle Triassic to Lower Jurassic continental deposits that provide essential information about the tetrapod assemblages of that time, documenting some of the oldest known dinosaurs and the first faunas numerically dominated by this group. The Upper Maleri Formation of the Pranhita-Godavari Basin preserves an early-middle Norian dinosaur assemblage that provides information about the early evolutionary history of this group in central-south Gondwana. This assemblage comprises sauropodomorph dinosaurs and an herrerasaurian, including two nominal species. Here, we describe in detail the anatomy of one of those early dinosaurs, the bagualosaurian sauropodomorph Jaklapallisaurus asymmetricus. The new anatomical information is used to investigate the position of the species in an updated quantitative phylogenetic analysis focused on early sauropodomorphs. The analysis recovered Jaklapallisaurus asymmetricus as a member of Unaysauridae, at the base of Plateosauria, together with Macrocollum itaquii and Unaysaurus tolentinoi from the early Norian of southern Brazil. This phylogenetic result indicates that the dispersal of early plateosaurian sauropodomorphs between the Southern Hemisphere and what nowadays is Europe would have occurred shortly after Ischigualastian times because of the extension of their ghost lineage. Thus, the presence of early plateosaurians in the early Norian of South America and India reduces a previously inferred diachrony between the biogeographic dispersals of theropods and sauropodomorphs during post-Ischigualastian times.

On a skeletally immature individual of Unaysaurus tolentinoi (Dinosauria: Sauropodomorpha) from the upper Triassic of southern Brazil.

The lineage of sauropodomorph dinosaurs raised some of the most impressive animals that ever walked on Earth. However, the massive titans of the Mesozoic Era originated from far smaller dinosaurs. The Triassic beds from Brazil yielded the earliest part of this evolutionary history. Despite the diverse fossil record of early sauropodomorphs, juvenile specimens, as well as certain species are poorly sampled. This is the case for Unaysaurus tolentinoi, an unaysaurid sauropodomorph from Caturrita Formation (ca. 225 Ma; early Norian, Late Triassic). The holotype and only specimen of U. tolentinoi was excavated from the Água Negra Locality (São Martinho da Serra, Rio Grande do Sul, Brazil) in 1998. More than two decades later, no other fossil vertebrates have been reported from the same fossiliferous site. Here we describe a skeletally immature specimen which was found in association with the holotype of U. tolentinoi. The specimen was discovered after a first-hand examination of the holotype and comprises some isolated vertebrae and elements from the posterior autopodium. According to linear regressions, its metatarsal I is approximately 41.7 mm in length, compared to approximately 75.9 mm in the holotype. The repeated elements and reduced size indicates that it does not belong to the elements originally used to erect U. tolentinoi. Rather, the specimen is assigned to U. tolentinoi by topotypy and shared morphology. In addition to the reduced size, distinct lines of evidence (e.g., neurocentral sutures; bone texture) support its assignment to a skeletally immature individual. In sum, the new material expands the record of U. tolentinoi, and represents an additional juvenile dinosaur from the Caturrita Formation.

The origin of an invasive air sac system in sauropodomorph dinosaurs.

One of the most remarkable features in sauropod dinosaurs relates to their pneumatized skeletons permeated by a bird-like air sac system. Many studies described the late evolution and diversification of this trait in mid to late Mesozoic forms but few focused on the origin of the invasive respiratory diverticula in sauropodomorphs. Fortunately, it is possible to solve this thanks to the boom of new species described in the last decade as well as the broad accessibility of new technologies. Here we analyze the unaysaurid sauropodomorph Macrocollum itaquii from the Late Triassic (early Norian) of southern Brazil using micro-computed tomography. We describe the chronologically oldest and phylogenetically earliest unambiguous evidence of an invasive air sac system in a dinosaur. Surprisingly, this species presented a unique pattern of pneumatization in non-sauropod sauropodomorphs, with pneumatic foramina in posterior cervical and anterior dorsal vertebrae. This suggests that patterns of pneumatization were not cladistically consistent prior to the arrival of Jurassic eusauropods. Additionally, we describe the protocamerae tissue, a new type of pneumatic tissue with properties of both camellae and camerae. This reverts the previous hypothesis which stated that the skeletal pneumatization first evolved into camarae, and derived into delicate trabecular arrangements. This tissue is evidence of thin camellate-like tissue developing into larger chambers. Finally, Macrocollum is an example of the gradual evolution of skeletal tissues responding to the fastly specializing Respiratory System of saurischian dinosaurs.

A giant phytosaur (Diapsida, Archosauria) from the Upper Triassic of India with new insights on phytosaur migration, endemism and extinction

AbstractA new Mystriosuchinae phytosaur,Colossosuchus techniensis, is described from the Upper Triassic Tiki Formation of India.Colossosuchusis diagnosed by multiple apomorphies, including a strongly downturned terminal rosette (c. 70°), closely spaced mediolateral band‐like ornamentation on dorsal surface of the nasal, dorsolaterally oriented supratemporal fenestra, ventrolaterally inclined postorbital–squamosal bars depressed below the skull table, dorsally convex parietal–squamosal bar that descends ventrolaterally below the skull roof, dome‐shaped skull in lateral view, prominent neural arch laminae and fossae, three sacral vertebrae, robust proximal end of the tibia, and fibula with high anteroposterior flaring of the distal end. The total body length of the largest individual recovered from the bonebed is estimated to be more than 8 m, suggesting thatColossosuchusis one of the largest phytosaurs known. Phylogenetic analysis nestsColossosuchusand other undescribed specimens from India within Mystriosuchinae. These form a distinct clade and represent the earliest record of endemism among Gondwanan phytosaurs. This clade is recovered as sister taxa to ((Volcanosuchus + Rutiodon) + Leptosuchomorpha), where the depressed supratemporal fenestra first appeared in the phytosaur lineage, a feature previously used to diagnose the derived leptosuchomorphs. Early‐diverging phytosaur diversification may have coincided with the final stages of the Carnian Pluvial Event with their possible migratory routes along the circum‐Tethyan coastline. The lineage continued to evolve mostly through endemic radiations and experienced an extinction event during early Norian, which marked the disappearance of most of the non‐leptosuchomorph taxa. This is attributed to post‐CPE aridification, although more study is required.

Provenance of the first terrigenous sediments in the western Sichuan Basin during the Late Triassic: Implications for basin evolution from marine to continental

The Late Triassic was a key period for the evolution of the western Sichuan Basin from marine to continental sedimentation. However, the provenance of the earliest terrigenous sediments during this period remains debated, hindering our understanding of the tectonic events that ruled the evolution of the basin at that time. Herein, samples of fine sandstone from the Upper Triassic Ma'antang and Xiaotangzi formations in the western Sichuan Basin were collected for petrology, heavy mineral analysis, bulk rock geochemistry, and detrital U–Pb dating. In addition, corresponding data from potential source areas were collected for comparison. The sedimentological, geochronological, and geochemical characteristics of terrigenous sediments suggest that the clastic materials were mainly sourced from the Qinling orogenic belt and Yangtze Craton (including the northern and western margin). The Longmen Shan thrust belt likely provided clastics since the Early Norian. Siliciclastic deposits of the Late Triassic sedimentary succession of the western Sichuan Basin (Ma'antang and Xiaotangzi formations) yielded young zircon U–Pb ages of 214–245 Ma, suggesting that these zircons were likely sourced from the magmatic activities in the South Qinling orogenic belt or Yidun Island Arc. Combined with previous research, this study predates the transformation of the western Sichuan Basin from marine to continental sedimentation in the Late Carnian/Early Norian period.

The elongated neck of sauropodomorph dinosaurs evolved gradually.

Discoveries from South America have increased our knowledge on the early evolutionary history of sauropodomorph dinosaurs. The dietary shift from faunivorous to herbivorous creatures and the increasing body size are both widely documented in the fossil record. Conversely, the initial evolution of the elongated neck is poorly known. It is one of the most diagnostic features of Sauropodomorpha. There is a gap between the record of short-necked sauropodomorphs from Carnian (±233 Ma) and long-necked forms from early Norian (±225 Ma). As a consequence, it is unknown if the cervical vertebrae became long gradually or abruptly. In the present study, we present a new specimen excavated from strata that belong to this time interval (±228 Ma). CAPPA/UFSM 0352 comprises a series of five cervical vertebrae unearthed from the Late Triassic of Southern Brazil. The vertebrae are proportionately longer than that of older forms and proportionately shorter than that of younger ones. Therefore, our results demonstrate that the elongation of the neck of sauropodomorphs is an example of gradual evolutionary process. Except by its elongated shape, the general anatomy of the cervical elements resembles that of the earliest forms (i.e., have a conservative anatomy). Combined with previous data, it is possible to conclude that the shape of the skull and teeth, as well as the neck proportions, were the first structures to clearly differ derived sauropodomorphs from early diverging forms. Finally, some of the recovered phylogenetic scenarios favor the origins of the elongated neck in the clade Bagualosauria.

Geomagnetic polarity and carbon isotopic stratigraphic assessment of the late Carnian -earliest Norian in Svalbard: evidence for a major hiatus and improved Boreal to Tethyan correlation

Geomagnetic polarity and carbon isotopic stratigraphic assessment of the late Carnian -earliest Norian in Svalbard: evidence for a major hiatus and improved Boreal to Tethyan correlation

An early-diverging crocodylomorph from the early Norian (Late Triassic) of Texas demonstrates a wide distribution of early members across low-latitude Pangea

An early-diverging crocodylomorph from the early Norian (Late Triassic) of Texas demonstrates a wide distribution of early members across low-latitude Pangea

Geochemical study on fossil vertebrates from some specific Permian and Triassic beds of the Paraná Basin (Brazil): A preliminary approach

Access to the geochemical conditions of paleoenvironments is made through the analysis of the chemical composition of the hydroxyapatite of fossil bones. This is because the bones were able to incorporate chemical elements from adjacent fluids into their pores and channels during the fossildiagenesis process, and as a result, have their original apatite substituted. The main goal of the present study was to seek geochemical signatures in bones and related sedimentary rocks from three well-known fossiliferous units of the Paraná Basin in southern Brazil: Irati Formation, Cisuralian (three bones/rocks sampled); Rio do Rasto Formation, Guadalupian (three bones/rocks sampled) and Santa Maria Supersequence, Late Ladinian–Early Norian (nine bones/rocks sampled). Fossils and rocks were analyzed by ICP-MS. Although the Permian fossil and rock samples (from Irati + Rio do Rasto Formations) have variations among them, some geochemical parameters are common (high Be–Co–Cu–Zn–Y–W–Pb contents). This composition differentiates them from the Triassic fossils and rocks (from Santa Maria Supersequence - high contents of V–As–Ba). The P content (essential in apatite) was used to determine the degree of preservation of the studied material. The better-preserved Permian specimen has a mean = 434.19 ratio (P content of fossil bones/P content of rock) and the lowest ratio = 1.87. Within the Triassic specimens, the better preserved has ratio = 205.64 and lowest = 2.24. In Permian specimens, Y shares similar ionic radio with Ca. For Triassic specimens, Y replaces Ca and As either replaces P or is associated with minerals in bone cavities. So, Y and As substitute Ca in the analyzed material. Therefore, the Permian samples are better preserved than the Triassic ones based on their P contents; Y and As can be used as indicators to differentiate Permian fossils (high Y values) from Triassic ones (high As values). These geochemical signatures can help in forensic cases, such as robbery or illegal transport of fossils.

Late Triassic rifting and volcanism on the northeastern Indian margin: A new phase of Neo-Tethyan seafloor spreading and its paleogeographic implications

The stratigraphic record of the northern Indian margin documents tectonic extension and magmatic activity during the Late Triassic. The geodynamic process and the paleogeographic implications of this event remain, however, incompletely understood. This study presents new stratigraphic, petrographic, geochronologic and geochemical data from a mixed carbonate-clastic Upper Triassic succession of the Tethys Himalaya in southern Tibet (Tela Formation). The new finding of volcanic detritus with bimodal mafic/felsic composition in the Tela Formation indicates nearby volcanic activity constrained by detrital zircon UPb dating as latest Carnian to early Norian in age (229–223 Ma). The alkalic signature of mafic volcanic rock fragments, showing LREE enrichment and HREE depletion, indicates a within-plate extensional setting. The igneous activity in northeastern India compares with coeval volcanism in northwestern Australia, suggesting that a rift-related volcanic belt developed along the northern margin of eastern Gondwana in the Late Triassic. This volcanic belt controlled the accumulation of thick turbidites of the Langjiexue Group in northeastern India and induced accelerated tectonic subsidence and increased sedimentation rates all along the Tethys Himalayan zone as far west as the Zanskar-Spiti Synclinorium. This major rifting phase is suggested to have initiated a new phase of sea-floor spreading in the Neo-Tethys Ocean that torn off the Lhasa Block from eastern Gondwana.

A new theropod dinosaur from a peculiar Late Triassic assemblage of southern Brazil

The early evolution of theropod dinosaurs is poorly understood. Whereas sauropodomorphs are well-known from the oldest dinosaur bearing outcrops, the record of theropods is fragmentary and ambiguous. The Triassic deposits from Brazil yielded some of the oldest dinosaurs worldwide. These dinosaurs came from two distinct Assemblage Zones (AZ): the Hyperodapedon AZ (Carnian) and the Riograndia AZ (early Norian). Here, a specimen previously assigned to cf. Dinosauromorpha is reassessed. CAPPA/UFSM 0157 comes from an enigmatic assemblage with predominance of the traversodotind cynodont Siriusgnathus. This assemblage has been tentatively assigned to the Riograndia AZ. However, the absence of index fossils still hampers a reliable assignation. The specimen, which comprises a proximal portion of a left femur, belongs to a new theropod taxon erected here as Erythrovenator jacuiensis gen. et sp. nov. The new dinosaur differs from all other known Triassic dinosaurs based on the absence of a raised dorsolateral trochanter of the femur. Erythrovenator jacuiensis gen. et sp. nov. is regarded as a theropod on the basis of the pyramidal shape of the anterior trochanter in anterior view. The results of a phylogenetic analysis corroborate this assignation. Therefore, the new dinosaur represents one of the oldest theropod dinosaurs worldwide, shedding lights on some of the earliest theropod features. Finally, the new specimen also represents the first carnivorous dinosaur from the assemblage dominated by the traversodontid cynodont Siriusgnathus, increasing our knowledge of the faunal content of this enigmatic assemblage.

Triassic faunal successions of the Paraná Basin, southern Brazil

The Paraná Basin was filled by a sedimentary package deposited in successive sedimentation episodes related to the tectonic events that hit the SW portion of the Gondwana. The Triassic portion of this package, known worldwide for its continental tetrapod fauna, occurs only in the southern portion of the basin and is represented by 2 s-order sequences: the Sanga do Cabral Supersequence (SCS - Early Triassic) and the Santa Maria Supersequence (SMS - Middle-Late Triassic). The SCS fauna, including temnospondyls, parareptiles (mainly Procolophon), archosauromorphs, putative synapsids, and a number of indeterminate specimens, is traditionally considered Early Triassic and corresponds to the “Procolophon abundant zone” of the Karoo Basin (the upper levels of the Lystrosaurus AZ), in the upper Katberg Formation, which is Induan to early Olenekian in age. The sedimentary environment of the SCS is thought to be a wide alluvial plain, in which small and shallow channels spread northwards into a vast semiarid environment. By its turn, the Middle-Upper Triassic Santa Maria Supersequence is divided into four third-order sequences, from base to top: Pinheiros-Chiniquá, Santa Cruz, Candelária and Mata. Each of these sequences begins with fluvial deposition (low sinuosity rivers) that is overlain by transgressive shallow lacustrine deposits. The first three of these sequences present a very rich record of fossil tetrapods, including four successive faunal associations: Dinodontosaurus Assemblage Zone (Ladinian, within the Pinheiros-Chiniquá Sequence), Santacruzodon AZ (Ladinian/Carnian, in the Santa Cruz Sequence), Hyperodapedon AZ (Carnian) and the Riograndia AZ (early Norian), the latter two respectively at the base and top of the Candelária Sequence. In general, the lower portion of the package (Pinheiros-Chiniquá and Santa Cruz Sequences) was deposited under more basic and dried environmental conditions and are dominated by synapsids, while the top of the section (Candelária Sequence) is characterized by more acid and humid conditions and by a shift in the faunal content, with diapsids as dominating forms and presenting an increase of diversity compared to the lower biozones.

The osteology of the early-diverging dinosaurDaemonosaurus chauliodus(Archosauria: Dinosauria) from theCoelophysisQuarry (Triassic: Rhaetian) of New Mexico and its relationships to other early dinosaurs

AbstractThe early evolution of dinosaurs is documented by abundant postcranial material, but cranial material is much rarer and comparisons of cranial features among early dinosaurs are limited to only a few specimens. Here, we fully detail the osteology of the unusual early-diverging dinosaur Daemonosaurus chauliodus from the latest Triassic Coelophysis Quarry in northern New Mexico, USA. The taxon possesses a unique and curious suite of character states present in a variety of early dinosaurs, and the morphology of D. chauliodus appears to link the morphology of Herrerasaurus with that of later diverging eusaurichians. Our phylogenetic analyses places D. chauliodus at the base of dinosaurs and our interpretation of the unusual mix of character states of D. chauliodus does not lead to a firm conclusion about its nearest relationships or its implications for the evolution of character state transitions at the base of Dinosauria. The combination of character states of D. chauliodus should not be ignored in future considerations of character evolution in early dinosaurs. As one of the last members of the earliest radiation of saurischians in the Carnian–early Norian, D. chauliodus demonstrates that members of the original diversification of dinosaurs survived until nearly the end of the Triassic Period.

The Late Triassic Ischigualasto Formation at Cerro Las Lajas (La Rioja, Argentina): fossil tetrapods, high-resolution chronostratigraphy, and faunal correlations

Present knowledge of Late Triassic tetrapod evolution, including the rise of dinosaurs, relies heavily on the fossil-rich continental deposits of South America, their precise depositional histories and correlations. We report on an extended succession of the Ischigualasto Formation exposed in the Hoyada del Cerro Las Lajas (La Rioja, Argentina), where more than 100 tetrapod fossils were newly collected, augmented by historical finds such as the ornithosuchid Venaticosuchus rusconii and the putative ornithischian Pisanosaurus mertii. Detailed lithostratigraphy combined with high-precision U–Pb geochronology from three intercalated tuffs are used to construct a robust Bayesian age model for the formation, constraining its deposition between 230.2 ± 1.9 Ma and 221.4 ± 1.2 Ma, and its fossil-bearing interval to 229.20 + 0.11/− 0.15–226.85 + 1.45/− 2.01 Ma. The latter is divided into a lower Hyperodapedon and an upper Teyumbaita biozones, based on the ranges of the eponymous rhynchosaurs, allowing biostratigraphic correlations to elsewhere in the Ischigualasto-Villa Unión Basin, as well as to the Paraná Basin in Brazil. The temporally calibrated Ischigualasto biostratigraphy suggests the persistence of rhynchosaur-dominated faunas into the earliest Norian. Our ca. 229 Ma age assignment to Pi. mertii partially fills the ghost lineage between younger ornithischian records and the oldest known saurischians at ca. 233 Ma.

Carnian–Norian (Late Triassic) climate change: Evidence from conodont oxygen isotope thermometry with implications for reef development and Wrangellian tectonics

The Carnian–Norian (C–N) transition (Late Triassic) has long been postulated as an interval of major climatic changes, though the nature of such changes and their ecological impact remains largely unexplored. We use oxygen isotopes measured on monogeneric conodont assemblages (δ18OPO4) from the Canadian Cordillera to trace seawater temperature evolution at the western margin of Pangea and in the allochthonous Wrangellia Terrane. Different conodont taxa show conspicuous offsets in δ18OPO4, suggesting that they had preferential habitat depths and genus-specific temperature corrections must be applied. Thus, δ18OPO4 from the Williston Lake sections indicates low mid-latitude sea surface temperatures (SSTs) ranging from ∼28 to 35°C, favouring a generally warm Late Triassic climate scenario. The parvus Subzone of C–N transition, which marks the peak faunal turnover, records major climatic perturbations: SSTs peaking at ∼34°C then decreasing to ∼29°C. This is followed by a quick temperature rebound and a second pulse of cooling from 33 to 27°C in the asymmetrica-Norigondolella Subzone, marking the coldest temperatures in the ∼20 Myr study interval. A secular warming trend towards the late Norian reinstated after the transient cooling in the earliest Norian, coinciding with reef expansions in the Late Triassic. The middle–late Norian (Alaunian 3–Sevatian 1) represents a hothouse climate comparable to the Paleocene–Eocene Thermal Maximum and its warmth supressed both diversity and origination rates of reef-building biota. Early Norian δ18OPO4 data from Frederick Island, Haida Gwaii (part of the Wrangellia Terrane) are more depleted in 18O compared to coeval samples from the Black Bear Ridge, indicating ∼3–5°C warmer temperatures than the Laurentian epicontinental sea. The temperature difference supports a more southern, probably sub-equatorial position for at least part of the Wrangellia Terrane during the earliest Norian.

The Foura Sandstone type section (Samaropollenites speciosus Zone, Carnian–early Norian; early Late Triassic), Timor-Leste: preliminary correlation between Timor and the Bonaparte Basin

The Foura Sandstone type section (<i>Samaropollenites speciosus</i> Zone, Carnian–early Norian; early Late Triassic), Timor-Leste: preliminary correlation between Timor and the Bonaparte Basin

Reassessment of Unaysaurus tolentinoi (Dinosauria: Sauropodomorpha) from the Late Triassic (early Norian) of Brazil, with a consideration of the evidence for monophyly within non-sauropodan sauropodomorphs

We present a systematic revision of the ‘prosauropod’ dinosaur Unaysaurus tolentinoi. Past phylogenetic treatments of this taxon have presented it as a key constituent of the low-diversity Plateosauridae. This clade, along with Massospondylidae, is currently regarded as a relatively non-controversial example of monophyly within non-sauropodan sauropodomorphs – an otherwise pectinate grade of animals from which the apically nested Sauropoda is derived. The inclusion of Unaysaurus within Plateosauridae is primarily dependent on similarities shared with the Northern Hemisphere taxon Plateosaurus, with a sister-taxon relationship between these taxa indicating the earliest known instance of Pangaean cosmopolitanism among early sauropodomorphs. However, an in-depth exploration of the character states that in the past have produced this relationship shows that many either present a complex distribution throughout early Sauropodomorpha, or are subject to ambiguities in the formulation and scoring of primary homologies. An updated cladistic analysis finds weak continued support for a sister-taxon relationship between Unaysaurus and Plateosaurus; however, this result is supported by synapomorphies that are either labile with respect to taxon inclusion, not exclusively restricted to the clade, or represented by character-state dichotomies of relatively subtle differentiation. These uncertainties aside, the validity of Unaysaurus tolentinoi remains supported by several unambiguous autapomorphies. Although strong anatomical evidence for monophyletic divergences amongst early sauropodomorphs remains elusive, the relatively low stratigraphical occurrence of Unaysaurus demonstrates that the typical ‘prosauropod’ bauplan – and hence habitual herbivory – had evolved by the early Norian.