Are Spondylosoma and Staurikosaurus (Santa Maria Formation, Middle-Upper Triassic, Brazil) the oldest saurischian dinosaurs?

Spondylosoma absconditum (Ladinian, Middle Triassic) lacks certain Dinosauriformes-Dinosauria characters (sigmoid curve and epipophyses in neck, distally placed deltopectoral crest on humerus) and those present (accessary hyposphene-hypantra vertebral articulations, three sacral vertebrae, elongate pubis) also occur in the Rauisuchia (Crurotarsi, Suchia, Rauisuchiformes).Spondylosoma is tentatively referred to the Rauisuchidae because, as indicated by the ventrolaterally directed sacral ribs, the ilium was held obliquely (and not vertically as in the Poposauridae).Staurikosaurus pricei (Carnian, Upper Triassic), one of the oldest dinosaurs, possesses autapomorphies of Dinosauriformes, Dinosauria, Saurischia, and Herrerasauridae, but it retains certain plesiomorphic characters (subcircular distal end to tibia, ?proximally placed deltopectoral crest, axially unshortened neural spine to sacral 1, ?scapular spine unreduced distally) not present inHerrerasaurus (Carnian, Ischigualasto Formation, Argentina). However, the femur is more derived in the loss of the trochanteric shelf and reduction of the lesser trochanter (as also occurs in some prosauropods). The two vertebrae sacrum ofStaurikosaurus is a plesiomorphic character for Dinosauria (also in other herrerasaurids, German Upper Triassic prosauropodSellosaurus gracilis — ?males), to which was added either a caudosacral (e.g. Santa Maria prosauropodSaturnalia, Sellosaurus gracilis — ?females) or a dorsosacral (e.g. Ischigualasto theropodEoraptor and prosauropodRiojasaurus). Agrosaurus (Upper Triassic, Durdham Down, England — not Australia) is not a herrerasaurid; it is referred to the prosauropod dinosaurThecodontosaurus antiquus (Upper Triassic, Durdham Down).     

Notes on the Melanorosauridae,a family of large Prosauropod Dinosaurs (Saurischia: Sauropodomorpha)

The femur of the anchisaurid (= plateosaurid) prosauropoddinosaur Euskelosaurus browniiHuxley, 1866 from the lower Elliot Formation (Upper Triassic) of South Africa is sigmoidal in posterior view with the fourth trochanter well removed from the medial edge. The sympatric Melanorosaurus readiHaughton, 1924 is not a junior synonym of Euskelosaurus brownii because the femur is straight in posterior view with the fourth trochanter close to the medial edge. In addition to Melanorosaurus, the Melanorosauridae Huene, 1929 includes Camelotia borealis nov. gen. and nov. sp. from the Upper Triassic of England, Riojasaurus incertusBonaparte, 1969 from the Upper Triassic of Argentina, and possibly some of the posteranial material referred to the? rauisuchian the codontian Sinosaurus triassicusYoung, 1948 from the Lower Jurassic of China.     

A Basal Sauropodomorph (Dinosauria: Saurischia) from the Ischigualasto Formation (Triassic,Carnian) and the Early Evolution of Sauropodomorpha

Background

The earliest dinosaurs are from the early Late Triassic (Carnian) of South America. By the Carnian the main clades Saurischia and Ornithischia were already established, and the presence of the most primitive known sauropodomorph Saturnalia suggests also that Saurischia had already diverged into Theropoda and Sauropodomorpha. Knowledge of Carnian sauropodomorphs has been restricted to this single species.

Methodology/Principal Findings

We describe a new small sauropodomorph dinosaur from the Ischigualsto Formation (Carnian) in northwest Argentina, Panphagia protos gen. et sp. nov., on the basis of a partial skeleton. The genus and species are characterized by an anteroposteriorly elongated fossa on the base of the anteroventral process of the nasal; wide lateral flange on the quadrate with a large foramen; deep groove on the lateral surface of the lower jaw surrounded by prominent dorsal and ventral ridges; bifurcated posteroventral process of the dentary; long retroarticular process transversally wider than the articular area for the quadrate; oval scars on the lateral surface of the posterior border of the centra of cervical vertebrae; distinct prominences on the neural arc of the anterior cervical vertebra; distal end of the scapular blade nearly three times wider than the neck; scapular blade with an expanded posterodistal corner; and medial lamina of brevis fossa twice as wide as the iliac spine.

Conclusions/Significance

We regard Panphagia as the most basal sauropodomorph, which shares the following apomorphies with Saturnalia and more derived sauropodomorphs: basally constricted crowns; lanceolate crowns; teeth of the anterior quarter of the dentary higher than the others; and short posterolateral flange of distal tibia. The presence of Panphagia at the base of the early Carnian Ischigualasto Formation suggests an earlier origin of Sauropodomorpha during the Middle Triassic.     

Partial hindlimb of Blikanasaurus cromptoni n. gen. and n. sp.,representing a new family of prosauropod dinosaurs from the upper triassic of South Africa

The holotype of the prosauropod dinosaur Blikanasaurus cromptoni n. gen. and n. sp., a partial hindlimb, is described from the lower Elliot Formation (= Red Beds) (upper Carnian or lower Norian; Upper Triassic) of Herschel district, Transkei, South Africa. The new prosauropod family Blikanasauridae is characterized by a hindlimb that is extremely stocky, especially the metatarsus, with the distal tarsals medially situated. Blikanasaurus was an early experiment in the direction of heavily-built, quadrupedal saurischians, but it was not on the evolutionary line that gave rise to the Sauropoda.     

Cyclotosaurus cf. PosthumusFraas (Capitosauridae,Stereospondyli) from the Huai Hin Lat Formation (Upper Triassic), Northeastern Thailand: With a note on capitosaurid biogeography

Part of a large capitosaurid skull, similar to that of Cyclotosaurus posthumus from the Upper Triassic of Germany, has been discovered in the upper part of the Huai Hin Lat Formation near Chulabhorn (Nam Phrom) Dam. This discovery is consistent with the presumed Norian age of this formation. Although the phylogeny of the Capitosauridae is still unclear, the group of Upper Triassic Cyclotosaurus species to which C. posthumus belongs is monophyletic and seems to be known only from Laurasia or Northwestern Gondwana (Morocco). The occurrence of C. cf. posthumus in Thailand is consistent with the hypothesis previously put forward, that this part of Southeast Asia was bound to Laurasia in Mesozoic times.     

New and little-known grylloblattids of the family Geinitziidae (Insecta: Grylloblattida) from the Triassic and Jurassic of Europe,Asia, and South Africa

The fossil history of the family Geinitziidae is reviewed. New taxa are described in the family: Shurabia hissarica, sp. nov. (Lower Jurassic of Tajikistan), Sh. shartegica, sp. nov. (Upper Jurassic of Mongolia), Sh. serrata, sp. nov. and Ginitzia sagulensis, sp. nov. (both Lower Jurassic of Kyrgyzstan). The subfamily Stegopterinae Sharov, 1961 is synonymized under Geinitziinae Handlirsch, 1906. The genera Minesedes Fujiyama, 1973 and Ominea Fujiyama, 1973 (Upper Triassic of Japan) are synonymized under Shurabia Martynov, 1937 and Geinitzia Handlirsch, 1906, respectively. Geinitzia varia Bode, 1953 (Lower Jurassic of Germany) and Fletchizia picturata Riek, 1976 (Upper Triassic of South Africa) are redescribed from their holotypes. F. kapokraalensis Wappler, 2001 and F. aleda Wappler, 2001 (both Upper Triassic of South Africa) are transferred to the genus Shurabia.     

The Kuehneotheriidae (Mammaliaformes) from Saint-Nicolas-de-Port (Upper Triassic,France): a Systematic Review

The origin and first diversification of mammals in the Upper Triassic remain poorly understood, in part because many fossil discoveries are not fully studied, and in part because the material remains poor. The Saint-Nicolas-de-Port quarry (Rhaetian, France) is the second most important locality that yielded remains of Kuehneotherium, after the fissure-fillings of the Glamorganshire (Lower Jurassic, Wales). This study identifies one new species of Kuehneotherium, K. stanislavi, sp. nov., and a new genus of Kuehneotheriidae, Fluctuodon necmergor, gen. et sp. nov. For these two new species, lower and upper molars are described and the first reconstructions of the postcanine row are proposed. Comparisons with material of Kuehneotherium from other Upper Triassic sites (Syren in Luxembourg, Emborough in England, and Jameson Land in Greenland) suggest two distinct Upper Triassic specific kuehneotheriid assemblages, respectively, west and east of the London Brabant Massif. They also suggest that the extinction event during the Triassic/Jurassic transition did not have a great impact on Kuehneotherium.     

Isopod crustacean fossils from the Cenomanian stratotype: five new species in suborders Cymothoida,Asellota and Valvifera

We report on a significant fossil collection of five isopod crustaceans that belong to five families from three suborders. These fossils were discovered in Le Mans (western part of Paris Basin, France) during the second half of the nineteenth century by the French paleontologist Edouard Guéranger. The historical quarry has been studied a few years before by the French paleontologist Alcide d’Orbigny and used as part of his Cenomanian stratotype (Leach, 1814). The collection consists of two species in the Cymothooidea Dana, 1852 (Cirolanidae Dana, 1852; Lantoceramiidae fam. nov.), two Valvifera G. O. Sars, 1883 (Chaetiliidae Dana, 1853; Idoteidae Samouelle, 1819) and one Asellota Latreille, 1802 (Stenetriidae Hansen, 1905), which were previously unknown from Upper Cretaceous strata as old as the Cenomanian age (roughly 93–99 ma). Although phylogenetic dating based on extant taxa has assigned Permian to Triassic age of origin for the Asellota, the Stenetriidae Hansen, 1905 specimen is the first fossil record for this family. Another asellotan, Fornicaris calligarisi Wilson and Selden, 2016, is known from the Triassic (Norian) dating from approximately 210–215 ma. The valviferans were unknown from Mesozoic strata, previously being found in the Oligocene of Romania and the Fur Formation, Upper Paleocene/Lowermost Eocene of Denmark (Polz, 2007). This diverse assemblage of fossils with taxa assignable to extant families and one new family provides evidence for the presence of a substantially modern isopod fauna as early as the Cenomanian.     

On the anatomy and relationships ofEfraasia diagnostica (Huene) n. gen., a prosauropod dinosaur (Reptilia: Saurischia) from the Upper Triassic of Germany

Palaeosaurus (?) diagnosticus Huene from the Stubensandstone (Upper Triassic, Norian) of Nordwürttemberg, Germany is made the type species of a new genus of prosauropod dinosaurEfraasia, which is referred to the family Anchisauridae. The anatomy ofEfraasia diagnostica is conservative and it was an ideal ancestor for the later and more specializedAnchisaurus polyzelus of North America. The slenderness of the manus and pes ofEfraasia andAnchisaurus is matched amongst prosauropods only by the feet ofGyposaurus capensis andThecodontosaurus antiquus. Staurikosaurus pricei Colbert from the Upper Triassic of Brazil is probably a very primitive theropod dinosaur.     

A bryozoan-like chaetetid (possible sclerosponge)from Jurassic-Cretaceous limestone near Orhaneli,Northwestern Turkey

Bryozoan-like fossils have been found rarely in Upper Jurassic or Lower Cretaceous limestone near Orhaneliin northwestern Turkey; they represent the chaetetid species Atrochaetetes alakirensisCUIF & FISCHER, 1974, possibly a sclerosponge, and previously known only from the lower part of the Upper Triassic.     

Paläobotanische untersuchungen in Nord-Iran. I. Nachweis nichtmariner obertrias am nordabfall des alburs-gebirges. 1. Grossform der pflanzenfunde von seltenen gattungen

New investigations in the Zirab-Karmozd area (northern Iran) by the author could testify that a lower part of the coal-bearing layers in this area belongs to the Upper Triassic, most probably to the Raetic. Species of the genera Scytophyllum Bornemann, Lepidopteris Schimper, Keraiaphyllum Frentzen (Sarmatiella Prinada), and Drepanozamites Harris are described. Evidence of a non-marine Upper Triassic in the Albourz Mountains is given for the first time.     

The prolacertid LepidosaurianCosesaurus avicepsEllenberger & Villalta, a claimed «Protoavian from the Middle Triassic of Spain

Cosesaurus aviceps, a small, probable diapsid reptile from the Ladinian (Middle Triassic) of Montral-Alcover (prov. Tarragona, Spain), has been reported as an ancestor of the Neornites. The only known specimen of this taxon is likely an inmature individual. The allometric growth models have been introduced in the character analysis, to infer the features of the adult Cosesaurus. For the present analysis, only the features that can be ascertained without doubt have been retained. The characters have been classified as primitive and derived. Cosesaurus shares all the ten derived characters we have found with Prolacertiformes, especially with Macrocnemus. The sister-group Macrocnemus + Cosesaurus is characterized by the relatively elongated neck in respect both to the skull and trunk length, 7–8 cervical vertebrae with the centra relatively elongated, and a relatively narrow distal femur region.     

Anatomy of the prosauropod dinosaurBlikanasaurus cromptoni (Upper Triassic, South Africa), with notes on the other tetrapods from the lower Elliot Formation

The holotype of the prosauropod dinosaurBlikanasaurus cromptoni Galton & van Heerden 1985, a partial hindlimb, is described from the lower Elliot Formation (Red Beds; Carnian, Upper Triassic) of the Eastern Cape, South Africa.Blikanosaurus, along with the sympatricEuskelosaurus andMelanorosaurus, probably represent the oldest record of the Prosauropoda (certainly so for articulated remains). The prosauropod family Blikanasauridae is characterized by a hind-limb that is extremely stocky, especially the metatarsus.Blikanasaurus was an early heavily-built, quadrupedal saurischian, but it was probably not the ancestor for the Sauropoda. The other tetrapods of the lower Elliot Formation are reviewed with particular emphasis on the carnivorous dinosaurAliwalia rex.     

Taxonomic study of the tetrahedral, pentagonal and hexagonal spongy spumellarian Radiolaria from the middle Carnian (Late Triassic) of the Köseyahya nappe (Elbistan, SE Turkey) and other Triassic localities

This article is a taxonomic study of all spongy radiolarian taxa with five to six coplanar and four tetrahedrally disposed spines or spongy arms occurring in the middle Carnian from the Köseyahya section, near the town of Elbistan, SE Turkey. This fauna is characteristic of the Tetraporobrachia haeckeli radiolarian Zone, and comes from an 8 m thick succession of clayey–cherty limestones occurring at the lower part of the section. In addition, a few species from the Middle and Upper Triassic from other areas have been also included in this study to improve some generic diagnoses and to better understand the diversity and evolutionary trends of some genera, subfamilies and families. The taxonomy at the generic and suprageneric levels is based primarily on the types of microsphere. This new approach allowed new taxonomic arrangements of genera and suprageneric units, and suggested new and unexpected phylogenetic relationships among these radiolarians and between them and younger radiolarians. The authors discuss and describe 42 species, of which 36 are new, and 12 genera of which three are new (Charlottalum, Pentaspongodisculus, and Trimiducaella). The genera Pseudohagiastrum Pessagno, Natraglia Pessagno, Cantalum Pessagno and others are reinterpreted. All the genera studied, except Charlottalum, are assigned to three subfamilies, of which two (Pseudohagiastrinae and Trimiducinae) are new, and to two families (Angulobracchiidae and Relindellidae). The genus Charlottalum is described to replace the genus Cantalum, which has been erroneously used until present for Late Triassic pantanelliids with four tetrahedrally disposed three-bladed spines.     

New beetles (Insecta,Coleoptera) from the Lower Triassic of European Russia

Fossil remains of beetles are described from two Lower Triassic localities: Entala (Induan) and Tikhvinskoe (Olenekian). Only one beetle fossil was previously known from the Lower Triassic of Tikhvinskoe. The fossils are rather few and poorly preserved, but they are worth describing as finds rare for the Lower Triassic. Five fossils from Entala most probably belong to beetles of the same species of the formal genus Pseudochrysomelites. Beetles of this genus are especially abundant in deposits close to the Permian–Triassic boundary and can be considered “disaster taxa.” There are no known cases, either in the Permian or in the Middle–Upper Triassic, of a random sample of five specimens belonging to a single species. This suggests that in the Entala oryctocenosis the species diversity of beetles is extremely low. All three beetle fossils found in Tikhvinskoe belong to beetles of different species, showing that diversity had already started to increase. However, it remained low, and all fossils belong to the formal family Schizocoleidae, and two of the three belong to the same genus, Pseudochrysomelites. The Khei-Yaga locality, which immediately follows Tikhvinskoe in time (topmost Olenekian or early Anisian), already contains beetles of the families Asiocoleidae and Permosynidae. In the Lower Anisian of the Buntsandstein, such typical Mesozoic beetles as Cupedidae and Coptoclavidae have been recorded. The appearance of such advanced beetles as early as the Lower Anisian suggests that the famous Permian–Triassic crisis was not as deep as it is usually believed, and many beetles survived it, disappearing, however, from the fossil record in the Early Triassic.     

Stratigraphic framework, discontinuity surfaces, and regional significance of Campanian slope to ramp carbonates from central Dalmatia, Croatia

The sedimentology, microfacies, and stratigraphic age (from planktonic and benthic foraminifera and strontium-isotope stratigraphy) of a 300-m-thick Upper Cretaceous carbonate succession from the Island of ?iovo (central Dalmatia, Croatia) were analyzed in order to determine the lithostratigraphic, depositional, and chronostratigraphic framework. The Cretaceous strata were deposited in the southern part of the long-lasting (Late Triassic to Paleogene) Adriatic-Dinaridic Carbonate Platform (ADCP), one of a few late Mesozoic, intra-Tethyan, peri-Adriatic (sub)tropical archipelagos. The succession is separated by a firmground formational boundary into two lithostratigraphic units: the underlying Middle to Upper Campanian Dol Formation consisting of slope pelagic limestone with intercalated turbidites and debrites, and the overlying Upper Campanian ?iovo Formation composed of outer-ramp bioclastic-lithoclastic and echinoderm-dominated packstone. Age, lithology, and depositional settings of the ?iovo Formation are different from other penecontemporaneous, regionally important inner-platform carbonate successions within the ADCP domain. Therefore, the ?iovo Formation is proposed here as a new lithostratigraphic unit. Regionally important condensed intervals in the form of at least two firmground surfaces, characterized by Thalassinoides burrows (with phosphatic mineralization) that belong to the Glossifungites ichnofacies, occur in the lowermost part of the ?iovo Formation. Abrupt shallowing of depositional environments at the boundary between the Dol and the ?iovo Formation, and the generation of the formational boundary firmground, likely correlate with the regionally recorded Upper Campanian Event that represents a global eustatic sea-level fall. A regionally important subaerial exposure surface with nodular calcrete, rhizoliths, and Microcodium aggregates in the upper part of the ?iovo Formation represents a regional subaerial unconformity that was recorded across the ADCP domain and was interpreted as a consequence of diachronous and differential uplift of various parts of the platform in response to the formation of a forebulge in front of the approaching Dinaridic orogen.     

Upper Triassic (Carnian) reef biota from the Sambosan Accretionary Complex, Kyushu, Japan

Calcified sponges, algae, and reef problematica are abundant yet poorly known from the Triassic of Japan. They are abundant in shallow-water carbonate, redeposited blocks of the Sambosan Accretionary Complex, Konosé Group, and southern Kyushu. Based on study of thin-sections from reef limestone exposed along the Kuma River, some important organisms and reef microfacies are described, which seem typical of Upper Triassic reef complexes. The most abundant reef organisms are hypercalcified sponges, including sphinctozoans, inozoans and chaetetids, followed by cyanophycean algae (including “Tubiphytes”-like organisms), and solenoporacean red algae. Loose sponge spicules in one thin-section also indicate the occurrence of rare hexactinellid sponges. Chambered demosponges described from the Konosé carbonate rocks include Solenolmia manon manon (Münster), Colospongia sp., Jablonskyia andrusovi (Jablonsky), several unidentified chambered sponges as well as the inozoid Permocorynella sp. 1 and Permocorynella sp. 2. Also present are chaetetid sponges and solenoporacean red algae belonging to Parachaetetes cassianus (Flügel) and Parachaetetes? sp. or Solenopora? sp. Especially abundant in thin-sections are cyanophyceans and “Tubiphytes”-like organisms. Among the organisms is Cladogirvanella Ott and Hedstroemia sp. The composition of the biota and presence of typical problematic organisms increases our knowledge of shallow-water Upper Triassic carbonate rocks in a remote setting in western Panthalassa. The composition of the biota indicates a mostly Carnian age. Most comparable organisms are known from both the northeastern and southern Tethys.     

New insects (Insecta: Eoblattida,Embiida) from the Permian of Russia and the Triassic of Kyrgyzstan,with observations on the origin of webspinners

Locustoblattina segmentata gen. et sp. nov. and L. marginata gen. et sp. nov., new members of the family Mesorthopteridae (Eoblattida) are described from the Middle Triassic of the Madygen locality (Ladinian Stage of the Kyrgyzstan). New members of the Alexarasniidae (Embiida), Alexarasnia limbata Aristov, sp. nov. and Nestorembia shcherbakovi sp. nov., are described from the Upper Permian Isady locality in the Vologda Region and the Madygen locality, respectively. Parastenaropodites fluxa Storozhenko, 1996 (Mesorthopteridae) is redescribed from Madygen. The origin of webspinners from within the Permian eoblattids and the composition of the superorder Blattidea are discussed.