Vertebrates from the Late Triassic Thecodontosaurus-bearing rocks of Durdham Down,Clifton (Bristol,UK)

Since the discovery of the basal sauropodomorph dinosaur Thecodontosaurus in the 1830s, the associated fauna from the Triassic fissures at Durdham Down (Bristol, UK) has not been investigated, largely because the quarries are built over. Other fissure sites around the Bristol Channel show that dinosaurs represented a minor part of the fauna of the Late Triassic archipelago. Here we present data on microvertebrates from the original Durdham Down fissure rocks, which considerably expand the taxonomic diversity of the island fauna, revealing that it was dominated by the sphenodontian Diphydontosaurus, and that archosauromorphs, including sphenosuchian crocodylomorphs, coelophysoid theropods, and the basal sauropodomorph Thecodontosaurus, were diverse. Importantly, a few fish teeth provide new information about the debated age of the fissure deposit, which is identified as lower Rhaetian. Thecodontosaurus had been assigned an age range over 20–25 Myr of the Late Triassic, so this narrower age determination (209.5–204 Myr) is important for studies of early dinosaurian evolution.     

The age and formation mechanisms of Late Triassic fissure deposits,Gloucestershire, England: Comments on Mussini,G. et al. (2020). Anatomy of a Late Triassic Bristol fissure: Tytherington fissure 2

In the Late Triassic the landscape NE of present-day Bristol, SW England was dominated by Carboniferous Limestone ridges and cuestas that became progressively buried by continental Mercia Mudstones and finally inundated during the Rhaetian marine transgression. Mussini et al. (2020) adopt the assertions of earlier collaborators back to Whiteside and Marshall (2008) that terrestrial vertebrate assemblages from sediments contained within karstic fissure systems in the former limestone ridges at Cromhall, Tytherington and elsewhere are restricted to the Rhaetian. We review and reject the sedimentological, stratigraphic, geomorphological and topographic arguments for this and reassert a long pre-Rhaetian (Norian) history for the vertebrate-bearing fissure systems at both Tytherington and Cromhall. We also reject the contemporaneous Rhaetian freshwater-seawater mixing zone dissolution model for the fissure systems adopted by Mussini et al. (2020) and reaffirm that the Tytherington and Cromhall fissures developed as conduit caves with a long Triassic history. Applying a new regional study of the Rhaetian transgressive surface, we also show that whilst the fissures at Cromhall remained sealed after the Norian, those at nearby Tytherington were re-exposed in the Late Rhaetian. Already partially filled with Norian sediments, the Tytherington fissures were subject to reworking on the seabed. Internal collapses, probably triggered by well documented repeated regional seismicity, led to the chaotic state of the Tytherington fills when downward moving Rhaetian marine components came to lie amongst and mix with earlier Norian terrestrial sediments. The vertebrate associations in the Tytherington fissures therefore contain a substantial Rhaetian input whilst those at Cromhall do not.     

The terrestrial fauna of the Late Triassic Pant-y-ffynnon Quarry fissures,South Wales,UK and a new species of Clevosaurus (Lepidosauria: Rhynchocephalia)

Pant-y-ffynnon Quarry in South Wales yielded a rich cache of fossils in the early 1950s, including articulated specimens of new species (the small sauropodomorph dinosaur Pantydraco caducus and the crocodylomorph Terrestrisuchus gracilis), but no substantial study of the wider fauna of the Pant-y-ffynnon fissure systems has been published. Here, our overview of existing specimens, a few described but mostly undescribed, as well as freshly processed material, provides a comprehensive picture of the Pant-y-ffynnon palaeo-island of the Late Triassic. This was an island with a relatively impoverished fauna dominated by small clevosaurs (rhynchocephalians), including a new species, Clevosaurus cambrica, described here from a partially articulated specimen and isolated bones. The new species has a dental morphology that is intermediate between the Late Triassic Clevosaurus hudsoni, from Cromhall Quarry to the east, and the younger C. convallis from Pant Quarry to the west, suggesting adaptive radiation of clevosaurs in the palaeo-archipelago. The larger reptiles on the palaeo-island do not exceed 1.5?m in length, including a small carnivorous crocodylomorph, Terrestrisuchus, and a possible example of insular dwarfism in the basal dinosaur Pantydraco.     

New haramiyidan and reptile fossils from a Rhaetian bedded sequence close to the famous ‘Microlestes’ Quarry of Holwell,UK

We describe here new Late Triassic haramiyidan mammaliaform and reptile fossils from near the classic ‘Microlestes’ Quarry’ at Holwell, Somerset, U.K., where Charles Moore discovered a huge collection of microvertebrates in the 1850s. Moore’s discoveries included the haramiyid Thomasia (formerly ‘Microlestes’ and Microcleptes) for which he achieved worldwide fame. Subsequently, despite much fossicking by researchers at Holwell, few new identifiable specimens of mammaliamorphs and lepidosaurs have been recorded and these were by Kühne in 1939. The new finds described here from a bedded sequence, not from a fissure, add significantly to our knowledge of the Holwell tetrapods and to the Rhaetian terrestrial faunas of the SW U.K. Our discovery of haramiyidan teeth includes a previously unknown type of Theroteinus, a genus not previously recorded from outside of the St-Nicholas-de-Port locality in France. An archosaur tooth, possibly from a phytosaur, is also recorded. The new lepidosaur specimens add significant detail to the recently described ‘basal’ rhynchocephalian Penegephyrosaurus curtiscoppi as well as demonstrating that the global diversity of Lepidosauria in the Late Triassic remains incompletely known.     

Dinosaur ichnofauna of the Upper Jurassic/Lower Cretaceous of the Paraná Basin (Brazil and Uruguay)

Upper Jurassic and Lower Cretaceous sedimentary layers are represented in the Brazilian Paraná Basin by the fluvio–aeolian Guará Formation and the Botucatu Formation palaeoerg, respectively, overlapped by the volcanic Serra Geral Formation. In Uruguay, the corresponding sedimentary units are named Batoví and Rivera Members (both from the Tacuarembó Formation), and the lava flows constitute the Arapey Formation (also in Paraná Basin). Despite the lack of body fossils in the mentioned Brazilian formations, Guará/Batoví dinosaur fauna is composed of theropod, ornithopod and wide–gauge sauropod tracks and isolated footprints, as well as theropod teeth. In turn, the Botucatu/Rivera dinosaur fauna is represented by theropod and ornithopod ichnofossils smaller than those from the underlying units. The analysis of these dinosaur ichnological records and comparisons with other global Mesozoic ichnofauna indicates that there is a size reduction in dinosaur fauna in the more arid Botucatu/Rivera environment, which is dominated by aeolian dunes. The absence of sauropod trackways in the Botucatu Sandstone fits with the increasingly arid conditions because it is difficult for heavy animals to walk on sandy dunes, as well as to obtain the required amount of food resources. This comparison between the Upper Jurassic and Lower Cretaceous dinosaur fauna in south Brazil and Uruguay demonstrates the influence of aridization on the size of animals occupying each habitat.     

Zeolites in fissures of granites and gneisses of the Central Alps

Six different Ca‐zeolite minerals are widespread in various assemblages in late fissures and fractures in granites and gneisses of the Swiss Alps. The zeolites formed as a result of water–rock interaction at relatively low temperatures (<250 °C) in the continental upper crust. The zeolites typically overgrow earlier minerals of the fissure assemblages, but zeolites also occur as monomineralic fissure fillings. They represent the youngest fissure minerals formed during uplift and exhumation of the Alpine orogen. A systematic study of zeolite samples showed that the majority of finds originate from three regions particularity rich in zeolite‐bearing fissures: (i) in the central and eastern part of the Aar‐ and Gotthard Massifs; (2) Gibelsbach/Fiesch, in a fissure breccia located at the boundary of Aar Massif and Permian sedimentary rocks; and (3) in Penninic gneisses of the Simano nappe at Arvigo (Val Calanca). Rail and road tunnel construction across the Aar‐ and Gotthard Massif provided excellent data on zeolite frequency in Alpine fissures. It was found that 32% (Gotthard NEAT rail base tunnel, Amsteg section) and 18% (Gotthard road tunnel) of all studied fissures are filled with zeolites. The number of different zeolites is limited to six species: laumontite, stilbite and scolecite are abundant and common, whereas heulandite, chabazite and epistilbite occur occasionally. Calcium is the dominant extra‐framework cation, with minor K and Na. Heulandite and chabazite contain Sr up to 29 and 10 mol.% extra‐framework cations respectively. Na and K contents in zeolites tend to increase during growth as a result of changes in fluid composition and/or temperature. The K enrichment of stilbite found in surface outcrops compared to subsurface samples may indicate late stage cation exchange with surface water. Texture data, relative age sequences derived from fissure assemblages and equilibrium calculations show that the Ca‐dominated zeolites precipitated from fluid with decreasing temperature in the order (old to young = hot to cold): scolecite, laumontite, heulandite, chabazite and stilbite. The necessary components for zeolite formation are derived from dissolving primary granite and gneiss minerals. The nature of these minerals depends, among other factors, on the metamorphic history of the host rock. Zeolites in the Aar Massif derived from the dissolution of epidote, secondary calcite and albite that were originally formed during Alpine greenschist metamorphism from primary granite and gneiss assemblages. Zeolite fissures occur in areas of H₂O‐dominated fluids. This is consistent with equilibrium calculations that predict a low CO₂ tolerance of zeolite assemblages, particularly at low temperature.     

A Rhaetian microvertebrate fauna from Stowey Quarry,Somerset, U.K.

Rhaetian bone beds have been described from many locations in south-west England, around Bristol. One site that has never been reported is Stowey Quarry, some 13?km south of Bristol. This quarry yielded Lias, and revealed thin Rhaetian units in the quarry floor, including two bone beds, the basal Rhaetian bone bed, and a second, higher bone bed, also within the Westbury Formation. The fossil fauna of both includes elements typical of Rhaetian bone beds elsewhere, but showing differences in faunal composition between the two. The basal bone bed yielded more specimens and more species, with three taxa (Sargodon tomicus, Rhomphaiodon minor and Hybodus cloacinus) exclusive to this bed and eleven identified in total. Severnichthys acuminatus accounts for more than 50% of the countable teeth from the basal bone bed, followed by Gyrolepis albertii with 20%, Lissodus minimus with 14% and Rhomphaiodon minor with 9%. The basal bone bed, as ever, is dominated by chondrichthyans, whereas the upper bone bed is dominated by osteichthyans, which form 90% of the non-dental remains. The only unique taxon is Dapedium, and Gyrolepis albertii is the most abundant species with 45% of countable teeth, followed by Severnichthys acuminatus with 38% and Duffinselache holwellensis with 11%. These faunal differences, and the equally good condition of specimens between both samples confirms that the upper bone bed is independent of the basal bone bed, and is not a reworked subsample.     

Dinosaur eggshells from the Santonian Milk River Formation of Alberta,Canada

The North American fossil record of dinosaur eggshells for the Cretaceous is primarily restricted to formations of the middle (Albian–Cenomanian) and uppermost (Campanian–Maastrichtian) stages, with a large gap in the record for intermediate stages. Here we describe a dinosaur eggshell assemblage from a formation that represents an intermediate and poorly fossiliferous stage of the Upper Cretaceous, the Santonian Milk River Formation of southern Alberta, Canada. The Milk River eggshell assemblage contains five eggshell taxa: Continuoolithus, Porituberoolithus, Prismatoolithus, Spheroolithus, and Triprismatoolithus. These ootaxa are most similar to those reported from younger Campanian–Maastrichtian formations of the northern Western Interior than they are to ootaxa reported from older middle Cretaceous formations (i.e., predominantly Macroelongatoolithus). Characteristics of the Milk River ootaxa indicate that they are ascribable to at least one ornithopod and four small theropod species. The taxonomic affinity of the eggshell assemblage is consistent with the dinosaur fauna known based on isolated teeth and fragmentary skeletal remains from the formation, although most ornithischians and large theropods are not represented by eggshell. Relative to the Milk River Formation eggshell, similar oospecies occurring in younger Cretaceous deposits tend to be somewhat thicker, which may reflect an increase in body size of various dinosaur lineages during the Late Cretaceous.     

The first record of a neonatal ornithopod dinosaur from Gondwana

Discrete post-embryonic teeth and bone fragments have been recovered from the matrix with the holotype skeleton (MPM‐10001) of the ornithopod dinosaur, Talenkauen santacrucensis Novas et al., 2004 (Upper Cretaceous, Argentina). The minute tooth crowns are 1 mm apicobasally tall and 1.7 mm mesodistally wide. The crowns are symmetrical and have a centrally located primary ridge on the lingual surface. Secondary ridges lead to five marginal denticles on both teeth. The tooth morphology is consistent with dentary teeth in euiguanodontids. There is no evidence of transport, suggesting that the material is autochthonous with respect to the adult body block of T. santacrucensis (MPM‐10001). Steeply inclined wear facets on the lingual surface and associated microstriae support the conclusion that the minute teeth were from a post-embryonic euiguanodontid dinosaur rather than early stage replacement teeth. The morphology, size, and wear of the teeth and small bone fragments found in the body block of MPM‐10001 suggest that this material belongs to a neonatal T. santacrucensis. This is the first record of neonatal ornithopod remains from Gondwana.     

The Bristol Dinosaur Project

Dinosaurs have been fascinating to the widest public since the 1840s, and that interest has grown step-wise ever since. Public interest has been harnessed over the years especially by museums in blockbuster exhibitions, and in the form of best-selling books and films. Here we describe a major educational initiative, the Bristol Dinosaur Project, which has run for ten years and has reached tens of thousands of children and adults, supported by substantial funding. The Bristol Dinosaur Project focuses on the fourth dinosaur ever named in the world, Thecodontosaurus, discovered in Bristol in 1834, and named in 1836. The dinosaur is not in itself spectacular, being only 1–2 m long, but its evolutionary role as one of the first plant-eating dinosaurs in the world justifies our current research, and provides a strong theme for the public presentation. Further, the fact that the dinosaur is found as disarticulated bones in ancient tropical cave systems, allows us to develop numerous key themes with all age groups: the geological time scale, continental drift, reconstruction of ancient environments, modern landscape analogues, the rock cycle, evolution, biomechanics, and critical assessment of geological and palaeontological evidence. These themes are of key importance for socio-economic and intellectual reasons, and yet are often poorly understood.     

Stratigraphy,biostratigraphy and C-isotopes of the Permian–Triassic non-marine sequence at Dalongkou and Lucaogou,Xinjiang Province,China

Measured lithostratigraphic sections of the classic Permian–Triassic non-marine transitional sequences covering the upper Quanzijie, Wutonggou, Guodikeng and lower Jiucaiyuan Formations at Dalongkou and Lucaogou, Xinjiang Province, China are presented. These measured sections form the framework and reference sections for a range of multi-disciplinary studies of the P–T transition in this large ancient lake basin, including palynostratigraphy, vertebrate biostratigraphy, chemostratigraphy and magnetostratigraphy. The 121 m thick Wutonggou Formation at Dalongkou includes 12 sandstone units ranging in thickness from 0.5 to 10.5 m that represent cyclical coarse terrigenous input to the lake basin during the Late Permian. The rhythmically-bedded, mudstone-dominated Guodikeng Formation is 197 m and 209 m thick on the north and south limbs of the Dalongkou anticline, respectively, and 129 m thick at Lucaogou. Based on limited palynological data, the Permian–Triassic boundary was previously placed approximately 50 m below the top of this formation at Dalongkou. This boundary does not coincide with any mappable lithologic unit, such as the basal sandstones of the overlying Jiucaiyuan Formation, assigned to the Early Triassic. The presence of multiple organic δ¹³C-isotope excursions, mutant pollen, and multiple algal and conchostracan blooms in this formation, together with Late Permian palynomorphs, suggests that the Guodikeng Formation records multiple climatic perturbation signals representing environmental stress during the late Permian mass extinction interval. The overlap between the vertebrates Dicynodon and Lystrosaurus in the upper part of this formation, and the occurrence of late Permian spores and the latest Permian to earliest Triassic megaspore Otynisporites eotriassicus is consistent with a latest Permian age for at least part of the Guodikeng Formation. Palynostratigrahic placement of the Permian–Triassic boundary in the Junggar Basin remains problematic because key miospore taxa, such as Aratrisporites spp. are not present. Palynomorphs from the Guodikeng are assigned to two assemblages; the youngest, from the upper 100 m of the formation (and the overlying Jiucaiyuan Formation), contains both typical Permian elements and distinctive taxa that elsewhere are known from the Early Triassic of Canada, Greenland, Norway, and Russia. The latter include spores assigned to Pechorosporites disertus, Lundbladispora foveota, Naumovaspora striata, Decussatisporites mulstrigatus and Leptolepidites jonkerii. While the presence of Devonian and Carboniferous spores and Early Permian pollen demonstrate reworking is occurring in the Guodikeng assemblages, the sometimes common occurrence of Scutasporites sp. cf. Scutasporites unicus, and other pollen, suggests that the Late Permian elements are in place, and that the upper assemblage derives from a genuine transitional flora of Early Triassic aspect. In the Junggar Basin, biostratigraphic data and magnetostratigraphic data indicate that the Permian–Triassic boundary (GSSP Level) is in the middle to upper Guodikeng Formation and perhaps as high as the formational contact with the overlying Jiucaiyuan Formation.     

A new ornithischian dinosaur and the terrestrial vertebrate fauna from a bone bed in the Wealden of Ardingly,West Sussex

The Wealden Supergroup of south-east England has long been of interest to palaeontologists because of its diverse flora and fauna. The Supergroup is Early Cretaceous in age, occupying the time period immediately after the enigmatic end-Jurassic extinction. Wealden faunas therefore have the potential to be informative about the tempo and mode of post-extinction recovery, but due to lack of exposure in this densely populated part of southern England, are difficult to sample. In the summer of 2012, a number of ex situ fossiliferous blocks of sandstone, siltstone and limestone were discovered from building excavations at Ardingly College, near Haywards Heath in West Sussex. The sedimentology of the blocks indicates that they are from the Valanginian Hastings Group, and that Ardingly College is underlain by the Grinstead Clay Formation, rather than the Ardingly Sandstone Member. The blocks contain a diverse invertebrate fauna and flora, as well as vertebrate remains, which are found in a distinct sandstone horizon that probably represents the Top Lower Tunbridge Wells pebble bed. A tooth from an ornithschian dinosaur cannot be referred to any of the ornithischian taxa known from the Wealden Supergroup, and therefore represents a new taxon. Teeth of the crocodilian Theriosuchus extend the known range of this taxon in the Wealden, while teeth of an ornithocheird pterosaur confirm the presence of these animals in the skies above the Wealden sub-basins. Fusainized plant remains and the wing-case of a cupedid beatle indicate that wildfire was a ubiquitous feature of the Weald Sub-basin during the Valanginian.     

Perinates of a new species of Iguanodon (Ornithischia: Ornithopoda) from the lower Barremian of Galve (Teruel,Spain)

Although Iguanodon is one of the most abundant and well-known of Europe's dinosaur genera, fossils of young specimens are very rare. Indeed, the fossil record contains very few examples of the young of any non-hadrosaurid iguanodontian. Here we report the discovery of 13 Iguanodon perinates from the Lower Cretaceous of Galve (Teruel, Spain). The characteristics of an adult and juvenile found nearby show these perinates to belong to a new species: Iguanodon galvensis sp. nov. The histological and osteological features of these young animals indicate them to have been in their first year of life. The taphonomic features of their remains, plus the finding of clearly embryonic vertebrae alongside them, suggest the perinates of this species remained in the vicinity of their nests for some time, possibly congregating in nursery areas.     

An abelisaurid from the latest Cretaceous (late Maastrichtian) of Morocco,North Africa

During the latest Cretaceous, distinct dinosaur faunas were found in Laurasia and Gondwana. Tyrannosaurids, hadrosaurids, and ceratopsians dominated in North America and Asia, while abelisaurids and titanosaurians dominated in South America, India, and Madagascar. Little is known about dinosaur faunas from the latest Cretaceous of Africa, however. Here, a new abelisaurid theropod, Chenanisaurus barbaricus, is described from the upper Maastrichtian phosphates of the Ouled Abdoun Basin in Morocco, North Africa on the basis of a partial dentary and isolated teeth. Chenanisaurus is both one of the largest abelisaurids, and one of the youngest known African dinosaurs. Along with previously reported titanosaurian remains, Chenanisaurus documents the persistence of a classic Gondwanan abelisaurid-titanosaurian fauna in mainland Africa until just prior to the end-Cretaceous mass extinction. The animal is unusual both in terms of its large size and the unusually short and robust jaw. Although it resembles South American carnotaurines in having a deep, bowed mandible, phylogenetic analysis suggests that Chenanisaurus may represent a lineage of abelisaurids that is distinct from those previously described from the latest Cretaceous of South America, Indo-Madagascar, and Europe, consistent with the hypothesis that the fragmentation of Gondwana led to the evolution of endemic dinosaur faunas during the Late Cretaceous.     

Calcium isotopes in fossil bones and teeth — Diagenetic versus biogenic origin

We present the first systematic study of Ca isotopes (δ^44/40Ca) in Late Triassic to Late Cretaceous dinosaur bones and teeth (enamel and dentin) from sympatric herbivorous and carnivorous dinosaurs. The samples derive from five different localities, and data from embedding sediments are also presented. Additional δ^44/40Ca in skeletal tissues from modern reptiles and birds (avian dinosaurs) were measured for comparison in order to examine whether the original Ca isotopic composition in dinosaur skeletal apatite was preserved or might have changed during the diagenesis and fossilization process.δ^44/40Ca of fossil skeletal tissues range from −1.62‰ (Tyrannosaurus rex enamel) to +1.08‰ (Brachiosaurus brancai bone), while values in modern archosaur bones and teeth range from −1.63‰ (caiman enamel) to −0.37‰ (ostrich bone). The average δ^44/40Ca of the three types of fossil skeletal tissue analyzed - bone, dentin and enamel - show some systematic differences: while δ^44/40Ca in bone exhibits the highest values, while δ^44/40Ca in enamel has the lowest values, and dentin δ^44/40Ca falls in between. Values of δ^44/40Ca in the remains of herbivorous dinosaurs (0.1-1.1‰) are generally higher than those of bones of modern mammalian herbivores (−2.6‰ to −0.8‰) and from modern herbivorous archosaurs, which exhibit intermediate δ^44/40Ca (−0.8‰ to −0.4‰). These systematic isotopic shifts may reflect physiological differences between dinosaurs, mammals and reptiles representing different taxonomic groups of vertebrates.Systematic offsets in skeletal apatite δ^44/40Ca between herbivorous and carnivorous dinosaurs are not obvious, indicating a lack of a clear-cut Trophic Level Effect (TLE) shift between herbivores and carnivores in dinosaurs. This observation can be explained if the carnivorous dinosaurs in this study fed mainly on soft tissues from their prey and did not ingest hard (calcified) tissue to much extent. The most striking indication that the primary δ^44/40Ca is actually preserved in most of the fossil teeth is a difference in δ^44/40Ca of about 0.35 ± 0.10‰ (1SD) between dentin and enamel, based upon 11 of 16 analyzed dentin-enamel pairs. This difference is close to that found in modern reptiles (0.28 ± 0.05‰), and strongly suggests that this tell-tale signature is a primary feature of the fossilized dinosaur material as well. Furthermore, simple mass balance calculations show that changes of the original δ^44/40Ca in bones and teeth by diagenetically-formed calcium-bearing minerals are either small or would require implausible high original δ^44/40Ca values in the skeletal apatite.     

The first lizard fossil (Reptilia: Squamata) from the Mesozoic of South Korea

Upper Cretaceous deposits in Mongolia, Chinese Inner Mongolia, and, more recently, southern China, have yielded individually rich and taxonomically diverser lizard assemblages. Here we describe the remains of a new terrestrial lizard, Asprosaurus bibongriensis gen. et sp. nov., from the Upper Cretaceous of South Korea. It represents the first record of a Mesozoic lizard from the Korean Peninsula and, although incomplete, is exceptional in its very large size. Characters of the mandible support attribution to crown-group Anguimorpha, with the closest similarities being to monstersaurs, the group represented today by the venomous North American Beaded lizard and Gila monster, genus Heloderma. This group is well-represented in the Upper Cretaceous fossil record in of eastern Asia, and the remains of large monstersaurs have been recovered from several dinosaur egg localities, suggesting dietary preferences similar to those of the living genus. The new Korean lizard, recovered from the Boseong Bibong-ri Dinosaur Egg Site, fits the same pattern.     

A new Triassic pterosaur from Switzerland (Central Austroalpine, Grisons), Raeticodactylus filisurensis gen. et sp. nov.

A new basal non-pterodactyloid pterosaur, Raeticodactylus filisurensis gen. et sp. nov., is reported. It has been discovered in shallow marine sediments from the Upper Triassic of the lowest Kössen beds (late Norian/early Rhaetian boundary) in the central Austroalpine of Canton Grisons (Switzerland). The disarticulated specimen is comprised of an almost complete skull and a partial postcranial skeleton. A high and thin bony, sagittal cranial crest characterizes the anterodorsal region of the skull. The large mandible, with an additional keel-like expansion at the front, partly matches the enlarged sagittal cranial crest. A direct and close relationship to Austriadactylus cristatus, the only known Triassic pterosaur with a bony cranial crest so far, cannot be established. The teeth of the premaxilla are monocuspid and exhibit very strongly bowed enamel wrinkles on the lingual side whereas the enamel is smooth on the labial side. These monocuspid teeth are large and fang-like. The numerous smaller teeth of the maxilla show three, four and five cusps. These are very similar to the teeth of the Triassic pterosaur Eudimorphodon ranzii. The humerus shows a thinner construction than that seen in other Triassic pterosaurs. The femur is quite unusual with a caput femoris perpendicular to the shaft. The bones of the extremities are almost twice as long as the ones from the largest Triassic specimen E. ranzii (MCSNB 2888). The newly described pterosaur is an adult, with a wingspan of approximately 135 cm. A morphofunctional analysis suggests that R. filisurensis was a highly specialized piscivore and possibly a skim-feeder.     

Newly Discovered Fish Faunas from the Early Triassic, Karoo Basin, South Africa, and their Correlative Implications

Recent collecting in exposures of the lowermost Burgersdorp Formation (Beaufort Group), of the Karoo Basin of South Africa, has revealed a previously unknown fish fauna from the Early Triassic (Scythian), lowermost Cynognathus Assemblage Zone (CAZ), which forms an important component of the total vertebrate assemblage. The newly discovered fish material includes lungfish, saurichthyids, and a large microfauna that includes numerous isolated chondrichthyan teeth, two fin spine fragments, and actinopterygian scales and teeth. The latest fish finds, together with the lowermost Cynognathus Assemblage Zone vertebrate faunas, make this Karoo Basin Assemblage Zone one of the most diverse Early Triassic faunal assemblages, comparable in faunal diversity to those from the Czatkowice Formation (Poland) and the Arcadia Formation (Australia). The presence of the lungfish Ptychoceratodus phillipsi in the early Middle Triassic Cynognathus Assemblage Zone (Subzone B), and in the underlying latest Early Triassic Cynognathus Assemblage Zone (Subzone A), indicates that these lungfish could serve as range index fossils within the CAZ, and thus are potentially useful biostratigraphic markers across the Early-Middle Triassic boundary. Furthermore the ‘new’ fish fauna provides a vital marine realm link in particular with the faunas of Madagascar and Australia, that is unavailable using the tetrapod faunal elements of the lower CAZ.