Halorina cryptica nov. ichnogen., nov. ichnosp., mass-occurrence of Upper Triassic crustacean microcoprolites from neptunian dikes and sills cutting the Dachstein-type carbonate platform and their paleoenvironmental significance (Northern Apuseni Mountains,Romania)

The new crustacean microcoprolite Halorina cryptica nov. ichnogen., nov. ichnosp., is reported from cryptic cavities cutting the Upper Triassic (Carnian to lower Rhaetian) Dachstein-type limestones from the Northern Apuseni Mountains, Romania. The new ichnotaxon is extremely abundant in cavities, neptunian dikes and sills filled with red ferruginous carbonate sediment. The associated microfauna consists of ostracods and rare foraminifera. The microfacies is represented by bioclastic coprolite-bearing wackestone-packstone to grainstone. The red ferruginous carbonate fillings are strongly bioturbated. The neptunian sill located at the top of the studied section contains a rich ichnofauna associated with brachiopod accumulations. It is dominated by the dimerelloid rhynchonellid Halorella, indicating a late Norian-early Rhaetian age. Although neptunian dikes and sills are rather common in the Dachstein-type carbonate platform that extended on the northwestern Tethyan margin, we report here the first record of mass-occurrence of crustacean microcoprolites in neptunian dikes and sills filled with red carbonate sediments of marine origin. They are interpreted here as cryptic cavities with specific palaeoenvironmental features (e.g., lack of light, abundant nutrients, intensive microbial activity) where crustacean arthropods thrived.     

Linkichnus Terebrans New Ichnogenus et Ichnospecies,an Insect Boring from the Late Triassic of the Germanic Basin,Southern Germany

The proposed new ichnotaxon Linckichnus terebrans n. ig. et isp., is a small test-tube shaped boring produced by insects in a sphenophyte stem and occurs in Upper Triassic (Keuper) terrestrial deposits.     

A palaeospinacid shark from the Upper Triassic of south-west England

The fin spines of a new neoselachian shark are described from die Upper Triassic of Aust Cliff, Avon and Holwell, near Frome in Somerset. The spines are assigned to Palaeospinax rhaeticus sp. nov. This record adds to the known diversity of Upper Triassic marine vertebrates from Britain and extends the range of the palaeospinacid sharks into the Rhaetian.     

Upper Triassic reefs of the Oman Mountains: Data from the South Tethyan margin

Summary The Upper Triassic reefal limestones of the Oman Mountains were investigated with respect to their microfacies, palaeontology and community structure. The reef fauna described and figured for the first time occurs in parautochthonous slope deposits of the Arabian platform (Sumeini Group) and in allochthonous reefal blocks (‘Oman Exotics’, Hawasina Complex). The ‘Oman Exotics’ are tectonically dislocated blocks, derived from isolated carbonate platforms on seamounts in the Hawasina basin or in the South Tethys Sea. The lithofacies and fauna of these blocks comprise a cyclic platform facies with megalodonts, reef and reef debris facies. The reefal limestones are dated as Norian/Rhaetian by benthic foraminiferal associations (Costifera, Siculocosta, Galeanella) and typical encrusting organisms (Alpinophragmium, Microtubus). Some small ‘Oman Exotics’ are of Carnian age. The shallow-marine organisms include scleractinian corals of different growth forms, ‘sphinctozoans’, ‘inozoans’ chaetetids, spongiomorphids, disjectoporids and solenoporacean algae as the main reef builders, various encrusters like microbes, foraminifers, sponges and many different problematical organisms for the stabilisation of the reef framework and a group of dwellers including benthic foraminifers, gastropods, bivalves and a few dasycladacean algae. The reef communities are characterized by the coverage of organisms and distributional pattern. Analogies with the coeval reef deposits from the European part of the Tethys have been recognized. Some species, now collected in Oman, were also reported from American and Asian localities.     

Trusheimichnus New Ichnogenus From the Middle Triassic of the Germanic Basin,Southern Germany

The study of Jurassic Zoophycos from south‐eastern France has led us to some original results concerning its morphological organization and paleoenvironmental significance. Zoophycos represents a spreite constituted by a coiling lamina, constructed upwards in sediments, with only one opening at the sediment‐water interface. Zoophycos, produced by a deep sediment feeder, is emplaced late in an almost coherent substrate, intermediate between soft and firm ground. This suggests periods of very low or no sedimentation during which time the trace was constructed.

The Zoophycos studied developed on slope areas, characterized by a particular style of sedimentation: periods of nutrient supplies coming from proximal areas, alternating with periods of sedimentary breaks. Other paleoecologic and paleoethologic characteristics of the Zoophycos‐producer are discussed here: working activity, competition behavior, substrate modification, growing pattern and burrow oxygenation are the most interesting.     

A new species of Leptocycas(Zamiaceae)from the Upper Triassic sediments of Liaoning Province,China

A new cycad,Leptocycas yangcaogouensis sp.nov.,was found in sediments from the Late Triassic in western Liaoning,China.The pinnately compound leaves(Pseudoctenis type)are screwed in a crown on the stem top.The leaflets are linear,with parallel veins and decurrent bases on the rachis.The leaf bases are persistent.The cataphylls intermix with the leaves.The female cone is ovoid in shape.The characteristics of the new plant are more similar to those of Leptocycas gracilis,a Triassic cycad from North America,but the new species differs from L.gracilis in the size of its stem(7-8 vs.3-5 cm in diameter,respectively),leaves(length × width 100 × 16 vs.30 × 7 cm,respectively)and leaf density along the stem(4-6 vs.1-2 bases/1 cm length,respectively).Both L.gracilis and L.yangcaogouensis,having leaves of the Pseudoctenis type,show a closer relationship to the extant Dioon of Zamiaceae.The present study provides evidence for the origin of the genus Dioon,which may have come from Leptocycas plants of the Triassic.It would be assumed that the extent cycads in Zamiaceae originate from the pteridosperms in the Late Paleozoic and have evolved through the stage of L.gracilis and L.yangcaogouensis in Late Triassic,and reaching the extant Dioon.     

A new species of Leptocycas (Zamiaceae) from the Upper Triassic sediments of Liaoning Province,China

A new cycad,Leptocycas yangcaogouensis sp.nov.,was found in sediments from the Late Triassic in western Liaoning,China.The pinnately compound leaves(Pseudoctenis type)are screwed in a crown on the stem top.The leaflets are linear,with parallel veins and decurrent bases on the rachis.The leaf bases are persistent.The cataphylls intermix with the leaves.The female cone is ovoid in shape.The characteristics of the new plant are more similar to those of Leptocycas gracilis,a Triassic cycad from North America,but the new species differs from L.gracilis in the size of its stem(7–8 vs.3–5 cm in diameter,respectively),leaves(length×width 100×16 vs.30×7 cm,respectively)and leaf density along the stem(4–6 vs.1–2 bases/1 cm length,respectively).Both L.gracilis and L.yangcaogouensis,having leaves of the Pseudoctenis type,show a closer relationship to the extant Dioon of Zamiaceae.The present study provides evidence for the origin of the genus Dioon,which may have come from Leptocycas plants of the Triassic.It would be assumed that the extent cycads in Zamiaceae originate from the pteridosperms in the Late Paleozoic and have evolved through the stage of L.gracilis and L.yangcaogouensis in Late Triassic,and reaching the extant Dioon.     

Swimming trace of a coelacanth fish from the Lower Keuper of south-west Germany

Parundichna schoelli igen. nov., isp. nov. from the Middle Triassic (Ladinian) Lower Keuper of Rot am See (Baden-Württemberg, Germany) consists of clusters of sigmoidal scratches symmetrically arranged in a plaited pattern. It is here interpreted as the swimming trace of a large coelacanth fish. In contrast to ichnospecies of Undichna there is no unpaired groove left by the tail fin. Instead, the sets of parallel scratches resulted from the pendulum motion of two pairs of appendages, the pectoral and pelvic fins, which acted in alternation, as in tetrapods. This strange mode of swimming is compared to films of modern Latimeria and to computer simulations; it probably corresponds to a particular foraging behaviour. The presence of fittingly-sized coelacanths is documented in coeval carbonates (Alberti-Bank) by disarticulated skeletal elements. Since the Lower Keuper represents a fluvial and estuarine facies of the receding Muschelkalk sea, we cannot be sure whether this trace was made in a marine or freshwater environment.     

An unusual new archosauriform from the Middle–Late Triassic of southern Brazil and the monophyly of Doswelliidae

Until now the Doswelliidae was considered a monospecific family including Doswellia kaltenbachi from the Late Triassic of North America. The phylogenetic position of this taxon remained enigmatic until recently, when a sister‐group relationship with the Proterochampsidae was suggested. In the present contribution we describe the new doswelliid species Archeopelta arborensis gen. et sp. nov. from the Middle–Late Triassic of Brazil. A cladistic analysis recovered Archeopelta, Doswellia, and Tarjadia within a monophyletic group of basal archosauriforms, the Doswelliidae. The monophyly of this family is supported by the presence of osteoderm ornamentation that is coarse, incised, and composed of regular pits and the presence of an unornamented anterior articular lamina. Archeopelta is more closely related to Doswellia than to other archosauriforms by the presence of basipterygoid processes anterolaterally orientated, dorsal centra with a convex surface, width of the neural arch plus ribs of the first primordial sacral that are three times the length of the neural arch, and iliac blade laterally deflected, with strongly convex dorsal margin, and a length less than three times its height. The phylogenetic analysis indicates that Doswellidae is the closest large monophyletic entity to Archosauria, which achieved a wide palaeolatitudinal distribution during the late Middle and Late Triassic time span. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 161 , 839–871.     

Microfacies and depositional environment of an Upper Triassic intra-platform carbonate basin: the Fatric Unit of the West Carpathians (Slovakia)

Facies associations of the Rhaetian Fatra Formation from the Veká Fatra Mts. (West Carpathians) were deposited in a storm-dominated, shallow, intra-platform basin with dominant carbonate deposition and variable onshore peritidal and subtidal deposits, with 21 microfacies types supported by a cluster analysis. The deposits are formed by bivalves, gastropods, brachiopods, echinoderms, corals, foraminifers and red algae, ooids, intraclasts and peloids. A typical feature is the considerable variation in horizontal direction. The relative abundance and state of preservation of components as well as the fabric and geometric criteria of deposits can be correlated with depth/water energy-related environmental gradients. Four facies associations corresponding to four types of depositional settings were distinguished: a) peritidal, b) shoreface, above fair-weather wave base (FWWB), c) shallow subtidal, above normal storm wave base and d) above maximum storm wave base. The depositional environment can be characterized as a mosaic of low-relief peritidal flats and islands, shoreface banks and bars, and shallow subtidal depressions. The distribution and preservation of components were mainly controlled by the position of base level (FWWB), storm activity and differences in carbonate production between settings. Poorly or moderately diverse level-bottom macrobenthic assemblages are dominated by molluscs and brachiopods. The main site of patch-reef/biostrome carbonate production was located below the fair-weather wave base. Patch-reef/biostrome assemblages are poorly diverse and dominated by the branched scleractinian coral Retiophyllia, forming locally dm-scale autochthonous aggregations or more commonly parautochthonous assemblages with evidence of storm-reworking and substantial bioerosion by microborings and boring bivalves.Facies types and assemblages are comparable in some aspects to those known from the Upper Triassic of the Eastern and Southern Alps (Hochalm member of the Kössen Formation or Calcare di Zu Formation), pointing to similar intra-platform depositional conditions. The absence of large-scale patch-reefs and poor diversity of level-bottom and patch-reef/biostrome assemblages with abundance of eurytopic taxa indicate high-stress/unstable ecological conditions and more restricted position of the Fatric intra-platform setting from the open ocean than the intra-platform habitats in the Eastern or Southern Alps.     

Upper Triassic Pacific vicariance as a test of geological theories

Upper Triassic vicariance that spans the Pacific involving terrestrial biotas in south east Asia/south western North America and Queensland/Chile/Argentina is sum- marized. These terrestrial and freshwater organisms did not migrate via high-latitude landbridges or across ocean barriers or Pangaea, are endemic to these vicariant fragments, and are mostly identical species. Rejoining the vicariant fragments is compatible with rapid earth expansion but is incompatible with other geological theories that call upon Panthalassa, Pacifica, displaced terranes and slow earth expansion. Vicariance biogeography yields a rigorous test of these models since its data are derived entirely independently of them. The Upper Triassic time-frame was selected because it immediately preceded the break-up of Pangaea.     

Trace fossils in fluvial deposits of the uppermost Stockton Formation (Late Triassic), Newark Basin,New Jersey

Abstract

Fluvial deposits of the uppermost Stockton Formation (Late Triassic), Newark Basin, west-central New Jersey have yielded an assemblage of trace fossils. Dominated by burrows, specimens include Cochlichnus anguineus, Helminthoidichnites tenuis, Planolites beverleyensis, Scoyenia gracilis, Spongeliomorpha carlsbergi, Treptichnus bifurcus, Treptichnus pollardi, plant remains, and an undetermined vertebrate trace fossil. The assemblage belongs to the Scoyenia ichnofacies. On the basis of stratification and primary sedimentary structures, the beds are interpreted as deposits in a meandering stream environment. Larval insects, wormlike forms, and arthropods are probably responsible for most of the animal traces in wet or moist channel, floodplain, and point bar sediments subject to subaerial exposure.     

The temnospondyl amphibian Cyclotosaurus from the Upper Triassic of Poland

Abstract:  A gap in the Late Triassic fossil record of the capitosaur amphibian Cyclotosaurus is filled by new material from lacustrine deposits at Krasiejów, Poland, corresponding in age to the Lehrberg Beds (late Carnian) of Germany. The skull of the Polish cyclotosaur is intermediate in several respects between that of Cyclotosaurus robustus from the middle Carnian Schilfsandstein of Germany and the younger C. mordax from the early Norian Stubensandstein. It shows a decrease in the width of the skull and in the degree of concavity of the posterior margin of the skull roof. The differences are significant enough to warrant erection of a novel species, the name Cyclotosaurus intermedius sp. nov. being proposed. The pectoral girdle, identified for the first time in Cyclotosaurus , suggests the genus was more fully adapted to an aquatic mode of life than was Paracyclotosaurus .     

Biostratigraphic significance of a new early sauropodomorph specimen from the Upper Triassic of southern Brazil

The Wachholz site (Caturrita Formation, Late Triassic), in Agudo, Rio Grande do Sul (RS), southern Brazil, has yielded several sauropodomorphs. This includes CAPPA/UFSM 0002, described here based on associated elements from the basalmost portion of the site. The specimen possesses a set of traits shared with typical ‘prosauropods’: a concave caudal margin of the trunk neural spines and a broadly convex proximal end of metacarpal V. However, it also retains some plesiomorphic features, for instance, the slender pedal digit I. Some bones closely resemble those of Unaysaurus tolentinoi, the other definitive sauropodomorph from the Caturrita Formation, an affinity corroborated by a new phylogenetic analysis. An updated biostratigraphic framework correlates the Wachholz, Água Negra (São Martinho da Serra/RS) and Botucaraí Hill (Candelária/RS) sites based on their sample of sauropodomorphs. In addition, the record of Jachaleria in the Botucaraí Hill site, a dicynodont also known from early Norian deposits of Argentina, indicates an equivalent age to deposits bearing U. tolentioni. Accordingly, a more constrained age is proposed for the Água Negra site. This is important as the early Norian marks the transition from an epoch of low sauropodomorphs representativeness to a period of extreme abundance of the group in Early-to-Middle Mesozoic ecosystems.     

Middle Triassic paleomagnetic data from northern Bulgaria: constraints on Tethyan magnetostratigraphy and paleogeography

Magnetostratigraphic and biostratigraphic data are presented from the Anisian (Middle Triassic) Peri-Tethyan Edivetur section of northwestern Bulgaria. A dual-polarity component of magnetization carried by magnetite delineates a magnetic stratigraphy of mainly reversed polarity. Magnetozones are dated by means of foraminifer and conodont biostratigraphy. Data from Edivetur are compared with data from Middle Triassic Tethyan limestone sections with the aim of contributing to the completion of the Middle Triassic magnetic polarity time scale. We also propose that paleomagnetic data from Edivetur can be used as proxy data for the paleogeographic position of the Moesian platform. The Moesian platform was located at 21–24°N along the southern margin of Europe. It was probably marginally separated, but not detached or rotated away from Europe by the North Dobrugea transtensional trough, which is interpreted as a back-arc basin resulting from the northward subduction of the Neo-Tethys (Vardar) or Paleo-Tethys ocean. Paleomagnetic data from this study and other minor tectonic elements are used to generate a paleogeographic sketch map of the Pangea-bounded western Tethys and Peri-Tethys at Middle/early Late Triassic time.     

Grallator-Dominated Fossil Footprint Assemblages and Associated Enigmatic Footprints from the Chinle Group (Upper Triassic), Gateway Area,Colorado

The upper part of the Chinle Group (Late Triassic) of the Gateway area in western Colorado is extraordinarily rich in fossil footprint assemblages. Dominant track types include small Grallator tracks, generally attributed to Coelophysis-like theropods, which often occur in high densities of 50 to 100 per m². Other abundant ichnotaxa that are attributable to dinosaurs or dinosaur-like archosaurs include Pseudotetrasauropus and Tetrasauropus, attributed to prosauropods and sauropods, respectively. Several Pseudotetrasauropus-like tracks appear to be functionally didactyl and may indicate a new ichnotaxon that represents an animal that shows certain unusual features that are convergent with dromeosaurs and certain birds. Such convergence may reflect inherent growth programs as much as functional adaptations. Non-dinosaurian ichnotaxa include Brachychirotherium (probably of aetosaur affinity) and Rhynchosauroides, attributed to a sphenodontid/lizard-like form. Other ichnotaxa include probable therapsid (dicynodont) tracks labeled Pentasauropus sp., mammaloid (non-therian mammal and/or mammal-like reptile) tracks, and the trails of arthropods. Excellent preservation and high track densities mark the Gateway assemblages in a thin stratigraphic interval in the upper part of the Chinle Group (Rock Point Formation). The track assemblages are similar to those reported from the Chinle Group in other parts of the Colorado Plateau and Rocky Mountain region, extending over most of Colorado, Utah, northern Arizona and northern and eastern New Mexico. Some of the Chinle ichnotaxa (Grallator and Brachychirotherium) are found in the overlying Wingate Formation, indicating that it is also Late Triassic in age, at least in the lower part. However, overall the Chinle and Wingate assemblages are quite different, most notably in the rarity of mammaloid/mammal-like tracks in the Chinle Group.     

First beetle elytra,abdomen (Coleoptera) and a mine trace from Lunz (Carnian,Late Triassic,Lunz‐am‐See,Austria) and their taphonomical and evolutionary aspects

Abstract: Four new species of Permosynidae are described: Platycrossus caroli Ponomarenko in Meller et al. sp. nov., Hydrobiites handlirschi Ponomarenko in Meller et al. sp. nov., Ademosyne polyzetete Ponomarenko in Meller et al. sp. nov. and Diarcuipenna bennettitophila Ponomarenko in Meller et al. sp. nov. Together with one polyphagous abdomen, they represent the first beetle remains from the Lunz formation, known for its richness in bennettite and cycad leaf remains and also for the bennettite reproductive organs. Furthermore, the first insect mine trace on a Nilssonia leaf segment is described, which is one of the most ancient linear mine traces. The rareness of insect remains in the plant‐bearing layers of the Lunz formation is still an enigma from the taphonomical–chemical point of view. The palaeoentomological and palaeobotanical considerations tentatively indicate a relationship between the rare occurrence of coniferous plants and the evolution or radiation of phytophagous insects during this time.     

A new species of the enigmatic archosauromorph Doswellia from the Upper Triassic Bluewater Creek Formation,New Mexico,USA

Abstract: Doswellia sixmilensis is a new species of the doswelliid archosauromorph genus Doswellia named for an incomplete skeleton from the Upper Triassic Bluewater Creek Formation of the Chinle Group in west‐central New Mexico, USA. D. sixmilensis differs from D. kaltenbachi Weems, the type and only other known species of Doswellia, in its larger size, higher tooth count and greater heterodonty, possession of keels on the cervical centra and the presence of discrete knobs or spikes on some osteoderms. The holotype of D. sixmilensis is the fourth occurrence of Doswellia and only the second occurrence of a Doswellia skull, which includes the previously unknown premaxilla and maxilla (and therefore the best dentition) and has the best‐preserved cervical vertebrae. Although it adds to our knowledge of the anatomy of Doswellia, this new information does not alter previous concepts of the phylogenetic relationships of the doswelliid genera, largely because they are so poorly known anatomically. The genus Doswellia is known from the Newark Supergroup in Virginia, and the Chinle Group in Texas, New Mexico and Utah, in strata of Otischalkian–Adamanian age. The type locality of D. sixmilensis is c. 43 m stratigraphically below a bed from which U‐Pb dating of detrital zircons yields a maximum depositional age of c. 220 Ma, so this is a reasonable approximate numerical age for D. sixmilensis.     

The first tunicate with a calcareous exoskeleton (Upper Triassic,northern Italy)

The first tunicates with a calcareous exoskeleton are reported from Late Triassic buildup‐slope deposits of the Dolomites. Although examples of this group have been known since the early 1900s from the middle–upper Permian of eastern Asia and Sicily as Khmeria, they were erroneously attributed to rugose corals. These early representatives are small, double‐valved, conical skeletons, which evolved into multi‐plated capsules with up to 35 opercula. The latter are joined along zigzag margins, which in life could probably be opened for the atrial and branchial siphons. The construction and shape of these skeletons distinguish them from plants or other invertebrate phyla, while they share several similarities with living tunicates, specifically to sessile ascidians. Apart from a soft‐bodied genus from the lower Cambrian of China, ascidians are known only from isolated spicules, which occur sporadically from the Lower Jurassic onwards. The calcareous skeleton of these Late Triassic tunicates consists of aragonitic fibres, which form spherulitic or clinogonal microstructures. It seems that the stellate aragonitic spicules of Jurassic to Recent ascidians are a vestige of Permian–Triassic ancestors, which after the Carnian lost the ability to construct compound solid skeletons but partly still retain a soft double‐valved or multi‐operculate cellulose‐like tunic. The following taxa are described as new: Order Khmeriamorpha with the genera Khmeria Mansuy and Zardinisoma gen. nov., and the following species: Khmeria stolonifera (late Permian), Khmeria minima (Late Triassic), Zardinisoma japonicum (late Permian), Z. cassianum, Z. pyriforme, Z. polyplacophorum and Z. pauciplacophorum (all Late Triassic).