Battenizyga, a new Early Triassic gastropod genus with a discussion of the caenogastropod evolution at the Permian/Triassic boundary

Battenizyga, a new Early Triassic gastropod genus from the Moenkopi Formation of Utah, is described and the speciesAnoptychia eotriassica Batten & Stokes, 1986 is placed in it. The new genus has an axially ribbed planktonic larval shell and a teleoconch with an angulated periphery. This character combination is unknown from the Palaeozoic. Therefore,Battenizyga represents additional evidence that recovery from the end-Permian mass extinction was connected with a faunal turnover. Additionally, the extinction of diverse Palaeozoic groups of the Caenogastropoda in the Permian (e.g., the Pseudozygopleuridae) suggest a turnover. All caenogastropod genera that hold Early Triassic species, have post-Palaeozoic type species and most were not reported from the Palaeozoic. This corroborates the view that there was an intense faunal turnover within the Caenogastropoda.Battenizyga is probably a caenogastropod that is closely related to the superfamily Zygopleuroidea which is abundant in the late Palaeozoic and early Mesozoic.      

Smithian (Early Triassic) ammonoids from Tulong, South Tibet

Intensive sampling of the Tulong Formation in South Tibet has facilitated the construction of a highly-resolved middle and late Smithian ammonoid succession. The new biostratigraphical sequence comprises the middle Smithian Brayardites compressus beds, Nammalites pilatoides beds, and the Nyalamites angustecostatus beds followed by the late Smithian Wasatchites distractus beds and Glyptophiceras sinuatum beds. This faunal succession correlates very well with that of other Tethyan sequences such as the Salt Range (Pakistan), Spiti (India), Oman and South China. The Smithian faunal sequence from Tulong contains several taxa with broad geographic distribution (e.g., Owenites, Paranannites spathi, Shigetaceras, Wasatchites), thus enabling correlation with faunal successions from areas outside the Tethys (e.g., USA, British Columbia, Arctic Canada, South Primorye, Siberia). Early Smithian ammonoid faunas are almost absent in Tulong because of a preservation bias (absence of carbonate rocks). Five new ammonoid genera (Brayardites, Nammalites, Nyalamites, Shigetaceras, Tulongites) and six new species (Brayardites crassus, Brayardites compressus, Prionites involutus, ?Subflemingites compressus, Tulongites xiaoqiaoi, Urdyceras tulongensis) are described.     

Early Triassic coprolites from Australia and their palaeobiological significance

Abstract:  Coprolites from the Arcadia Formation, Queensland, Australia, were studied in conjunction with the vertebrate fossil assemblages from two localities to maximize our understanding of the palaeoecology of these Early Triassic deposits. Criteria used by other researchers to identify the producers of coprolites were found to be of little value in the Arcadia Formation specimens. Using a combination of shape, biostratigraphic distribution, size and included remains some of the coprolites are attributed to basal archosauromorphs and fish whereas others could not be identified. Perhaps the most important attribute of the Arcadia coprolites is that they preserved rare organisms such as cyanobacteria, insects and other arthropods, and a diversity of fish. Estimates of the number of actinopterygians and dipnoans preserved in coprolites significantly increased relative abundance estimates based on skeletal elements alone. Although coprolites are an important source of palaeobiological information, this information is limited by our poor understanding of the taphonomic processes involved in the fossilization of faecal matter and by the near impossibility of assigning coprolites to specific producers.     

The First Record of the Pterosaur Ichnogenus Purbeckopus in the Late Berriasian (Early Cretaceous) of Northwest Germany

A hypichnium of a manus imprint (preserved as plaster cast) indicates for the first time the presence of the large pterosaur ichnotaxon Purbeckopus cf. pentadactylus Delair, 1963 in the late Berriasian of northwest Germany. It is only the second record of Purbeckopus globally and the first pterosaur track from Germany. It provides evidence of a very large pterosaur (wingspan c. 6 m) in this area and from this time period not yet represented by skeletal remains. When compared with the English type material, the specimen exhibits some differences that are related mostly to different properties of the substrate on which both were left. These include, in the German track, an impression of the metacarpo-phalangeal joint of the wing finger, normally not present in pterosaur tracks. Also interesting is the rather blunt termination of the deeply impressed digits I–III, indicating rather short and blunt claws, which seem more suitable for walking than for grasping or climbing. The specimens of Purbeckopus in England and Germany occur in different environments: the English locality was situated close to a brackish lagoon, while the German site belongs to a limnic-deltaic system at the margin of a large, freshwater lake.     

The biogeography of Early Triassic ammonoid faunas: Clusters, gradients, and networks

The aim of this paper is to quantitatively investigate the spatial and temporal biogeographical relationships of the recovery of ammonoid faunas after the Permian-Triassic mass extinction using three complementary numerical approaches among which is a new, non-hierarchical clustering strategy. The faunal data set consists of a taxonomically homogenised compilation of the spatial and temporal occurrences of ammonoid genera within 20 Early Triassic Tethyan and Panthalassic sites ranging from 40°S to 70°N in palaeolatitudes. In addition to hierarchical cluster analysis (hCA) and nonmetric multidimensional scaling (NMDS), we introduce a third, new non-hierarchical clustering technique allowing the visualisation of a nonmetric interassemblages similarity structure as a connected network constructed without inferring additional internal nodes. The resulting network, which we call a “Bootstrapped Spanning Network” (BSN), allows the simultaneous identification of partially or totally nested as well as gradational linear or reticulated biogeographical structures.The identified interlocalities relationships indicate that the very beginning of the Early Triassic (Griesbachian) corresponds to a very simple biogeographical context, representing a time of great cosmopolitanism for ammonoids. This context shifts rapidly to a more complex configuration indicative of a more endemic and latitudinally-restricted distribution of the ammonoids during the middle and late Early Triassic (Smithian and Spathian). From an evolutionary dynamic point of view, our results illustrate a very rapid (less than ca. 1.4 myr) Early Triassic recovery of the ammonoid faunas, in contrast to many other marine organisms. This recovery is linked with a marked increase in the overall biogeographical heterogeneity, and parallels the formation of a latitudinal gradient of taxonomic richness, which may be essentially controlled by the progressive intensification of the gradient of sea surface temperature. From a methodological point of view, we show that a BSN is a simple, intuitively legible picture of the nested as well as gradational taxonomic similarity relationships, hence providing a good synthesis (and additional insights) between hierarchical clustering and ordination in reduced space results.     

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.     

Early Triassic bivalves from the Feixianguan Formation in Xingyi,Guizhou and the Ximatang Formation in Qiubei,Yunnan (southern China)

Based on newly collected materials from the Lower Triassic Feixianguan Formation of Xiongwu section in Xingyi, Guizhou and the Ximatang Formation of Ximatang section in Qiubei, Yunnan, southern China, nine species belonging to seven genera are described. Two bivalve assemblages are recognized and regionally correlated in South China. The bivalve assemblage from the Feixianguan Formation of Xiongwu exhibits a higher diversity including seven species belonging to seven genera: Claraia griesbachi, Leptochondria virgalensis, Entolium (Entolium) microtis, Towapteria scythicum, Bakevillia exporrecta, Bositra sp., and Unionites? fassaensis. In contrast, the bivalve assemblage from the Ximatang Formation of Ximatang has a much lower diversity, consisting of only two species, i.e., Claraia griesbachi and Claraia radialis. Additionally, C. griesbachi and C. radialis, as the most common species of Claraia in the Early Triassic, are revised.     

Palaeoecological significance of a new Griesbachian (Early Triassic) gastropod assemblage from Oman

A new Early Triassic (Griesbachian) gastropod fauna from the Al Jil Formation of Oman is described. Early Triassic faunas from elsewhere (e.g. the Italian Dolomites and the western USA) are typically of low diversity and high dominance, usually attributed to environmental stress in the immediate aftermath of the end-Permian mass extinction event. The new Oman fauna has, by contrast, relatively high diversity, low dominance and a more even spread of individuals between taxa. It is the most diverse Griesbachian fauna known to date. This is attributed to the favourable (i.e. well-oxygenated) conditions under which the fauna lived. This uncharacteristic Griesbachian gastropod fauna demonstrates that, in the absence of oceanic anoxia, biotic recovery after the end-Permian extinction event may occur surprisingly quickly (within one conodont zone). The fauna is also partially silicified, which has increased its preservation potential relative to other Griesbachian gastropod assemblages. However, only one reappearing Lazarus genus is present in the Oman fauna. This suggests that there was some other control on the abundance of Lazarus genera at this time, other than the absence of silicified faunas as previously suggested.     

Ammonoid age control of the Early Triassic marine reptiles from Chaohu (South China)

Here we described a series of ammonoid specimens from the Early Triassic of Chaohu, South China and recognized the occurrence of Procolumbites for the first time in this area. The Procolumbites layer is about one meter above the first appearance of Chaohusaurus, indicating that the oldest Chaohusaurus is within the Procolumbites Zone of middle Spathian age. This new age constrain is significantly older than the previously suggested Subcolumbites Zone assignment (early late Spathian). To date, Chaohusaurus is the oldest known ichthyopterygian.     

Chondrichthyan teeth from the Early Triassic Paris Biota (Bear Lake County,Idaho, USA)

A new, diverse and complex Early Triassic assemblage was recently discovered west of the town of Paris, Idaho (Bear Lake County), USA. This assemblage has been coined the Paris Biota. Dated earliest Spathian (i.e., early late Olenekian), the Paris Biota provides further evidence that the biotic recovery from the end-Permian mass extinction was well underway ca. 1.3 million years after the event. This assemblage includes mainly invertebrates, but also vertebrate remains such as ichthyoliths (isolated skeletal remains of fishes). Here we describe first fossils of Chondrichthyes (cartilaginous fishes) from the Paris Biota. The material is composed of isolated teeth (mostly grinding teeth) preserved on two slabs and representing two distinct taxa. Due to incomplete preservation and morphological differences to known taxa, the chondrichthyans from the Paris Biota are provisionally kept in open nomenclature, as Hybodontiformes gen. et sp. indet. A and Hybodontiformes gen. et sp. indet. B, respectively. The present study adds a new occurrence to the chondrichthyan fossil record of the marine Early Triassic western USA Basin, from where other isolated teeth (Omanoselache, other Hybodontiformes) as well as fin spines of Nemacanthus (Neoselachii) and Pyknotylacanthus (Ctenachanthoidea) and denticles have been described previously.     

The Late Triassic Phytosaur Mystriosuchus Westphali, With A Revision of the Genus

Mystriosuchus westphali is based on a large, well-preserved cranium and a snout fragment from the Stubensandstein (Norian) of south-west Germany. The hypodigm is redescribed and new or poorly known cranial structures in phytosaurs are discussed. For the first time, the presence of a premaxillary crest is substantiated in a phytosaur. The type specimen shows a supernumerary occipital element (='tabular') that is probably fused to the parietal in other phytosaurs, and an orbitosphenoid. A computerised parsimony analysis confirms the hypothesis that Mystriosuchus is nested within Pseudopalatinae, the most derived clade of phytosaurs, and thus does not fall within basal phytosaurs. Mystriosuchus is characterised by five unique features (slit-like interpremaxillary fossa, triangular cross-section of the postorbito-squamosal bar, strongly reduced posttemporal fenestra, and two features of the cranial sculpture), plus eight synapomorphies that also occur in some more distantly related taxa. Mystriosuchus westphali is diagnosed by, among other apomorphies, a distinct premaxillary crest, a squamosal-proo¨tic contact, absence of a posterior process of the squamosal, and a slit-like posttemporal fenestra. The type species Mystriosuchus planirostris shows, most significantly, the naris facing forward anteriorly and upward posteriorly, and the longest rostrum and the highest degree of depression of the supratemporal opening in any phytosaur. Mystriosuchus exemplifies a common pattern in phytosaurids in being a genus that includes a gracile, elongated, slender-snouted and a more robust species with a broader, often crested snout. This study demonstrates that a detailed analysis of the cranial anatomy and the rigorous application of cladistic principles to identified character states help to clarify current inconsistencies in the taxonomy and nomenclature of phytosaurs.     

Endocranial anatomy and life habits of the Early Triassic archosauriform Proterosuchus fergusi

Proterosuchids are an important group of carnivorous basal archosauriforms characterized by a bizarre and enigmatic downturned premaxilla that overhangs the lower jaw. They are particularly significant because they radiated in the immediate aftermath of the Permian–Triassic mass extinction and represent one of the best known ‘disaster taxa’ following that event. While traditionally considered to be semi-aquatic, recent histological studies and geological data have suggested that it is more likely that they inhabited terrestrial environments. By using computed tomographic (CT) data, we virtually reconstruct the brain endocast and endosseous labyrinths of two adult specimens of Proterosuchus fergusi from the earliest Triassic of South Africa, in an attempt to understand its life habits within the context of basal archosauriform evolution. Endocasts reveal that the brain cavity is tubular in shape and the endosseous labyrinths are highly pyramidal. The angle of the lateral semicircular canal suggests that P. fergusi naturally held its head upwards ~17°, while the length of the cochlear duct suggests its auditory abilities were specialized towards low-frequency sounds. Furthermore, beam theory analysis suggests that the rostrum of P. fergusi is highly resistant to both bending and torsion when compared to modern crocodilians, although this resistance is neither enhanced nor reduced by the overhanging premaxilla. Comparative anatomical analyses suggest P. fergusi was probably a semi-aquatic, generalist apex predator capable of surviving the harsh environmental perturbations of the Early Triassic.     

Extremely distal fluvial sandstone within the playa system of Arnstadt Formation (Norian,Late Triassic), Central Germany

The Wachsenburg Sandstone of Thuringia (Central Germany) occurs within playa deposits of the Arnstadt Formation (Late Triassic, Norian) and furnishes an example of ephemeral river metamorphosis under dryland conditions. Characterized by high flow regime features, the sand-dominated lithofacies constitution exhibits sedimentation by channel processes under the influence of recurring flash floods. Bearing signatures of subaerial exposures, the fining-upward lithofacies cycles are bound by low-angle lateral accretion elements suggesting deposition in a meandering stream. Channel migration in response to point bar expansion and active bank erosion, led to the development of four laterally shifting point bar events. Unimodal palaeocurrent patterns with low variance and azimuthal dispersion support the point bar origin of the Wachsenburg Sandstone. With reduced water budget under largely semi-arid climate, the river progressively became smaller, highly sinuous and ultimately abandoned. The resulting point bar succession was finally covered with sheet flow deposits of over bank origin. The sandstone was deposited during a period of low base-level when the playa system temporarily fell dry.     

High-resolution biochronology and diversity dynamics of the Early Triassic ammonoid recovery: The Smithian faunas of the Northern Indian Margin

Based on new collections of abundant and well preserved material from the Salt Range (Pakistan), Spiti (Northern India) and Tulong (South Tibet), several recent studies focused on the taxonomic revision and detailed biostratigraphy of Smithian ammonoids. In this work, biochronological data for these three well-documented basins are analyzed by means of the Unitary Associations method, resulting in a biochronological scheme of unprecedented high-resolution for the Smithian of the Northern Indian Margin (NIM). Data for each basin are first processed separately, thus yielding three local biochronological zonations. Then, the three sequences are processed together as a regional three-section data set for the construction of an inter-basin sequence at the NIM level. The latter zonation comprises 16 Unitary Associations grouped into 13 zones for the entire Smithian. Analysis of ammonoid diversity dynamics based on this new highly resolved time frame highlights (i) a marked diversification during the early Smithian, (ii) a severe extinction during the late Smithian, and (iii) an overall very high turnover throughout the Smithian. At a global spatial scale and stage resolution, the diversity of Smithian ammonoid genera appears surprisingly high, as highlighted by a previous study. It is shown that at a smaller geographic scale and with the most highly resolved time frame, Smithian ammonoids of the NIM reached their explosive diversity peak essentially through extremely high turnover rates rather than through a classic diversification process of high origination rates coupled with low extinction rates. Based on recently published U/Pb ages, regional apparent total rates of origination and extinction of more than 100 species per My can be inferred for the Smithian ammonoids of the NIM.     

Plants of Leptostrobus Heer （Czekanowskiales） from the Early Cretaceous and Late Triassic of China, with Discussion of the Genus

Investigation of the Mesozoic seed plant Leptostrobus Heer from the Yangcaogou Formation of the Late Triassic and the Yixian Formation of the Early Cretaceous, Liaoning Province, China, provides new Insight Into Its general morphology and geographical distribution. The materials of L. cancer from the Yixian Formation described herein are later than all the past findings of this species and add to the record of L. cancer during the Early Cretaceous. Based on well-preserved specimens, the specific diagnosis Is slightly emended and the reconstructlon of L. cancer Is perfected. The materials from the Yangcaogou Formation of the Late Triassic are placed in L. spheericus, in addition, we review the history of investigation of the genus Leptostrobus since its establishment in 1876 and discuss the main characteristics of each species.     

Deciphering the exceptional preservation of the Early Triassic Paris Biota (Bear Lake County,Idaho, USA)

After the end-Permian mass extinction, the Early Triassic (∼251.9 to 247 Ma) is characterized by several biotic crises that particularly affected marine faunas; accordingly, marine ecosystems from this unstable interval have been often described as heavily depauperate. This assumption, however, may relate to a biased fossil record. The discovery of taphonomic windows, like Konservat-Lagerstätten, in the Early Triassic would help to better understand the composition and diversity of ecosystems at that time. The Paris Biota (Idaho, USA) is a highly diverse fossil assemblage from the earliest Spathian (early late Olenekian, ∼250.6 Ma), indicating a rapid rediversification for many groups after the end-Permian crisis and pointing toward a remarkably complex marine ecosystem ∼1.3 m.y. after the Permian-Triassic boundary. However, its detailed taphonomy has not yet been investigated. Here we present the mineral characterization of four of its most abundant taxa: discinoid and linguloid brachiopods, leptomitid sponges, and caridean shrimps. For this purpose, we combined data from Raman microspectroscopy, Fourier Transform InfraRed spectroscopy, and SEM-EDXS. Although all taxa were preserved in calcium phosphate, the morphology, structuring and size of crystals are highly dissimilar at a nano- to micrometric scale. In brachiopods, the ultrastructure of calcium phosphate shows unorganized bacillary-like crystals, while in crustaceans their size is considerably smaller and round-shaped. Similar small crystals are observed in sponges. However, the ultrastructure of calcium phosphate in sponges exhibits a well-defined preferential orientation. In addition, sponges show some compressed but preserved three-dimensional features, with an inner surface better preserved. Such analyses are essential to understand the taphonomic pathways enabling exceptional preservation. The further comprehension of preservation features would help to understand potential bias on observed diversity signals and their interpretation.     

Early Miocene Amphibians (Caudata, Salientia) from the Mokrá-Western Quarry (Czech Republic) with comments on the evolution of Early Miocene amphibian assemblages in Central Europe

The Mokrá-Western Quarry exhibits the rare occurrence of Early Miocene (MN 4) vertebrate fauna within the area of the eastern part of Central Europe. In addition to a rich fauna of reptiles and mammals, two fossiliferous karst joints (Mokrá-Western Quarry, 1/2001 Turtle Joint and Mokrá-Western Quarry, 2/2003 Reptile Joint) yielded a rich fauna of amphibians including 13 amphibian taxa: Salamandridae: Mioproteus sp., Chelotriton sp., type I, Chelotriton sp., type II, Triturus aff. roehrsi, Triturus cf. marmoratus, Triturus sp. (T. cristatus species group), Chioglossa meini, Mertensiella mera, Salamandridae gen. and sp. indet.; Pelobatidae: Pelobates sanchizi; Ranidae: Rana sp. (synklepton Rana esculenta); Bufonidae: Bufo sp. The first records of the West European species Triturus cf. marmoratus and Chioglossa meini are reported from the eastern part of Central Europe indicating the wide distribution of those taxa throughout the whole of Europe as early as MN 4. The oldest known record of Pelobates sanchizi documents the Early Miocene presence of representatives closely related to the extinct Late Oligocene representatives of Pelobates. The slow evolution of amphibian species is documented by the presence of Triturus cf. marmoratus and the oldest known occurrence of the extinct salamander Mertensiella mera.     

Latest Smithian (Early Triassic) ammonoid assemblages in Utah (western USA basin) and their implications for regional biostratigraphy,biogeography and placement of the Smithian/Spathian boundary

The late Smithian extinction represents a major event within the Early Triassic. This event generally corresponds to a succession of two, possibly three successively less diverse, cosmopolitan ammonoid assemblages, which when present, provide a robust biostratigraphic framework and precise correlations at different spatial scales. In the western USA basin, known occurrences of latest Smithian taxa are rare and until now, have only been documented from northeastern Nevada. Based on these restricted basinal occurrences, a regional zone representing the latest Smithian was postulated but not corroborated, as representative taxa had not yet been reported from outside Nevada. Here we document two new ammonoid assemblages from distant localities in northern Utah, overlying the late Smithian Anasibirites beds and characterized by the unambiguous co-occurrence of Xenoceltites subevolutus and Pseudosageceras augustum. The existence of a latest Smithian zone in the western USA basin is therefore validated, facilitating the identification of the Smithian/Spathian boundary and intra-basin correlation. This zone also correlates with the latest Smithian zone recognized from southern Tethyan basins. Additionally, these new data support other observed occurrences of Xenoceltites subevolutus throughout most of the late Smithian.