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The new neopterygian fish taxon Luoxiongichthys hyperdorsalis gen. et sp. nov. is established on the basis of five specimens from the second member of the Guanling Formation (Anisian, Middle Triassic) from Daaozi Quarry, Luoping, Yunnan Province, Southwest China. The new taxon is characterized by the following characters: triangular body outline with a distinct apex located between skull and dorsal fin; free maxilla; slender preopercular almost vertical; three suborbitals; at least eight strong branchiostegals with tubercles and comb−like ornamentation on the anterior margin; clavicles present; two postcleithra; ganoid scales covered by tubercles and pectinate ornamentation on the posterior margin with peg−and−socket structure; hemiheterocercal tail slightly forked. Comparison with basal actinopterygians reveals that the new taxon has parasemionotid−like triangular symplectics, but a semionotid opercular system. Cladistic analysis suggests that this new genus is a holostean, and either a basal halecomorph or basal semionotiform.
Traces of burrowing organisms from Lower Muschelkalk carbonate sediments of the Holy Cross Mountains (Góry Świętokrzyskie) interpreted as burrow systems of enteropneusts, have been described. Morphological and palaeoecological analysis of Triassic forms based on the comparison with the burrows of Recent enteropneusts is given. The presence of many horizons with burrows of enteropneusts in the profiles of the Lower Muschelkalk deposits (Łukowa beds) and the lithological characters of these deposits seem to indicate that the sedimentation took place in a zone of the basin affected by the activity of tidal currents.
A newly discovered silicified brachiopod interval from the Upper Member of the Guanling Formation (Late Anisian, Middle Triassic) in Guizhou Province (South China) is described for the first time. The most remarkable feature of this brachiopod assemblage, besides the very good preservation, is the very low taxonomic evenness and diversity. This impoverished, low diversity/high density assemblage is represented by more than 700 recovered specimens belonging to three species within two spiriferinid genera (Pseudospiriferina multicostata, P. pinguis, and Punctospirella fragilis). It is characterized by the overwhelming abundance of an endemic spiriferinid species, P. multicostata, which contributes to more than 90% of the community. Silicified valves of P. multicostata and Punctospirella fragilis allow detailed descriptions of the internal morphology based on direct observation. Brachiopod paleoecology, assessed by considering host−rock lithology, shell disarticulation, and shell size suggests that this endemic brachiopod fauna represents a favourable niche for development of dense brachiopod−dominated communities, i.e., high energy, hard substrate, nutrient rich environment.
Furcophyllia is an unusual coral with septa regularly splitting into branching sets called septal brooms. This pattern of septal apparatus is so alien to scleractinians, that, despite a trabecular microstructure of septa resembling that of the Scleractinia, the genus was originally ascribed to a rare group of corals informally referred to as sleractiniamorphs, previously known from the Ordovician and Permian. Genus Furcophyllia emerged together with corals of several groups, after the post−Permian crisis diversification of skeletonized anthozoans, some of them markedly differing in their skeletal features from typical Scleractinia. So far, the genus was represented by middle Carnian Furcophyllia septafindens from the Dolomites, in the Southern Alps. Here, we report Furcophyllia shaitanica sp. nov. from limestone boulders found in the volcano−clastic deposits of the upper Ladinian Šajtan suite of the South Eastern Pamirs. A new species of Furcophyllia signifies that the genus was a faunal element widely distributed in the Tethys.
The Bravaisberget Formation in Spitsbergen embraces an organic carbon-rich, clastic sequence that reflects a general shallow shelf development of the Middle Triassic depositional system in Svalbard. New observations and measurements of the type section of the formation at Bravaisberget in western Nathorst Land allow to present detailed litho- stratigraphical subdivision of the formation, and aid to reconstruct its depositional history. The subdivision of the formation (209 m thick at type section) into the Passhatten, Somovbreen, and Van Keulenfjorden members is sustained after Mørk et al. (1999), though with new position of the boundary between the Passhatten and Somovbreen mbs. The Passhatten Mb is defined to embrace the black shale-dominated sequence that forms the lower and middle parts of the formation (160 m thick). The Somovbreen Mb (20 m thick) is confined to the overlying, calcite-cemented sequence of marine sandstones. The Van Keulenfjorden Mb (29 m thick) forms the topmost part of the formation composed of siliceous and dolomitic sandstones. The formation is subdivided into twelve informal units, out of which eight is defined in the Passhatten Mb (units 1 to 8), two in the Somovbreen Mb (units 9 and 10), and also two in the Van Keulenfjorden Mb (units 11 and 12). Units 1, 3,5,7 and 9 contain noticeable to abundant phosphorite, and are interspaced by four black shale sequences (units 2,4,6, and 8). Unit 9 passes upwards gradually into the main sandstone sequence (unit 10) of the Somovbreen Mb. The base of the Van Keulenfjorden Mb is a discontinuity surface covered by thin phosphorite lag. The Van Keulenfjorden Mb consists of two superimposed sandstone units (units 11 and 12) that form indistinct coarsening-upward sequences. The Bravaisberget Fm records two consequent transgressive pulses that introduced high biological productivity conditions to the shelf basin. The Passhatten Mb shows pronounced repetition of sediment types resulting from interplay between organic-prone, fine-grained environments, and clastic bar environments that focused phosphogenesis. The lower part of the member (units 1 to 5) contains well-developed bar top sequences with abundant nodular phosphorite, which are under- and overlain by the bar side sequences grading into silt- to mud-shale. The upper part of the member (units 6 to 8) is dominated by mud-shale, showing the bar top to side sequence with recurrent phosphatic grainstones in its middle part. Maximum stagnation and deep-water conditions occurred during deposition of the topmost shale sequence (unit 8). Rapid shallowing trend terminated organic-rich environments of the Passhatten Mb, and was associated with enhanced phosphogenesis at base of the Somovbreen Mb (unit 9). Bioturbated sandstones of the Somovbreen Mb (unit 10) record progradation of shallow-marine clastic environments. The sequence of the Van Keulenfjorden Mb (units 11 and 12) was deposited in brackish environments reflecting closure of the Middle Triassic basin in western Svalbard.
Several fern remains are described from the para-autochthonous early late Ladinian flora of the Monte Agnello (Dolomites, N-Italy). The plants are preserved in subaerially deposited pyroclastic layers. Some ferns, known already from the Anisian and Ladinian of this area, are confirmed (Neuropteridium elegans), but several taxa are described for the first time (Phlebopteris fiemmensis sp. nov., Cladophlebis ladinica sp. nov., Chiropteris monteagnellii sp. nov.). Cladophlebis sp. and some indeterminable fern remains cannot yet be assigned to any family. Phlebopteris fiemmensis is now the oldest formally established species in the genus. The fern family Dipteridaceae (Thaumatopteris sp. and some fragments probably belonging to the Dipteridaceae because of their venation) has not been recorded previously from European sediments as old as the Ladinian. Although stratigraphically attributed to the late Ladinian, the flora is markedly distinct from other Ladinian floras of the Dolomites and the Germanic Basin. The flora from Monte Agnello shows a higher diversity in ferns than coeval floras from this area although characteristic elements of the Ladinian of the Dolomites such as Anomopteris and Gordonopteris are missing. The new flora misses also the Marattiales (e.g., Danaeopsis, Asterotheca) and other elements such as Sphenopteris schoenleiniana, typical for the Ladinian of the Germanic Basin.
Capitosaurs were the largest and homogeneous group of Triassic temnospondyl amphibians with cosmopolitan distribution. However, their interrelationships are debated. The first capitosaur cranial remains found in the Iberian Peninsula were assigned to Parotosuchus; herein, a re−description of this material, together with information on other remains recovered from the same site, enables us to classify them as a new genus: Calmasuchus acri gen. et sp. nov. (Amphibia: Temnospondyli) from the early−to−middle Anisian (early Middle Triassic). This capitosaur had a combination of plesiomorphic and non−plesiomorphic characters, such as posterolaterally directed tabular horns, paired anterior palatal vacuities, and unique morphology of the lower jaw. By cladistic analysis, we propose a new phylogeny for the monophyletic capitosaurs. In the analysis, Capitosauria is supported by seven synapomorphies. Wetlugasaurus is the most basal member of the clade. The score of the Russian taxon Vladlenosaurus alexeyevi resulted in a clade including Odenwaldia and the latter taxa. The Madagascarian Edingerella is the sister taxon of Watsonisuchus. Finally, Calmasuchus acri, the new taxon described here, appears as a more derived form than Parotosuchus. The new genus is the sister taxon of the Cyclotosaurus–Tatrasuchus and Eryosuchus–Mastodonsaurus clades.
A new genus Sinosaurichthys of the Saurichthyidae with three new species, S. longipectoralis, S. longimedialis, and S. minuta, are described and compared with Saurichthys. The new genus is represented by more than a hundred almost complete skeletons, collected from the strata corresponding to the Upper Member of the Guanling Formation (Pelsonian, Anisian, Middle Triassic) of two localities: Yangjuan of Panxian County, Guizhou Province, and Dawazi of Luoping, Yunnan Province, China. Sinosaurichthys differs from Saurichthys in having an unusual dermal pectoral girdle, high insertion of pectoral fin, relatively dorsally positioned axial skeleton in the abdominal region, and in the absence of branchiostegal rays. These differences are supposed to reflect the different life styles between the two genera. Sinosaurichthys, like the extant needlefish, probably has a better ability for cruising in surface water than Saurichthys. In addition, these species of Sinosaurichthys are compared, and their morphological differences also probably reflect individual adaptations for different habitats at the two localities.
The fossil record of coelacanths is patchy, with very few taxa known from the Triassic of Asia. We report here two new genera and species of coelacanths from the Luoping Biota, a recently found site of exceptional fossil preservation from Yunnan, South China. The first new taxon, Luopingcoelacanthus eurylacrimalis, is based on four specimens, which to− gether show most aspects of the anatomy. One specimen shows two small coelacanths inside the ventral portion of the ab− dominal cavity, and these are interpreted as intrauterine embryos, close to birth size, based on comparisons with previ− ously reported embryos of the fossil coelacanths Rhabdoderma and Undina, and the extant genus Latimeria. Our new find extends the evidence for ovoviviparity in coelacanths back from the Late Jurassic to the Middle Triassic. The second new taxon, Yunnancoelacanthus acrotuberculatus, is based on one specimen, and differs from Luopingcoelacanthus in the dentary, lachrymojugal, number of rays of the first dorsal fin, and especially in the ornament on dermal bones and scales. Acladistic analysis shows that the new taxa are closest relatives to the derived clade Latimerioidei. The relatively high di− versity of coelacanths in the Early Triassic, and adaptations of living Latimeria to low−oxygen conditions, suggests that the group may have included ‘disaster taxa’ that benefited from anoxic and dysoxic ocean conditions in the aftermath of the end−Permian mass extinction.
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Basal Archosauriformes had a wide geographic distribution through the Lower to Middle Triassic. Osmolskina czatkowiensis gen. et sp. nov. from Early Olenekian karst deposits at Czatkowice, west of Cracow, provides the first record from Poland. The reconstructed skull and attributed postcranial elements show a morphology closely resembling that of the Early Anisian African genus Euparkeria Broom, 1913, while differing at generic level. Both genera display the same mosaic of plesiomorphic and apomorphic character states, but share no unique apomorphic character state. They might thus be combined in the family Euparkeriidae Huene, 1920, but could also constitute two plesions of the same grade lying just below the Archosauria + Proterochampsidae node. Currently, Euparkeriidae remains monotypic because no other genus can be assigned to it with confidence. Until this problem is resolved, the term “euparkeriid” essentially denotes a grade of Lower to Middle Triassic non−archosaurian archosauriforms that are more derived than proterosuchid grade taxa, but lack the specializations of either erythrosuchids or proterochampsids. They were probably Pangaean in their distribution.
New examinations of numerous steinkerns of the Middle Triassic nautiloid Germanonautilusfrom southern Germany revealed new anatomic, ecologic, and taphonomic details, which are compared with Recent Nautilus. The attachment structures of the cephalic retractor muscle (large scar) and of the dorsal (black layer) and the posterior mantle (posterior narrow scar, anterior band scar of the mantle and septal myoadhesive bands), some with tracking bands (recording the anteriorward movement of the soft body during ontogeny), were seen in several specimens. The shape and proportions of these soft−tissue attachment structures resemble those of Recent Nautilus macromphalus and indicate a similar soft part anatomy. Based on their conch geometry, the mode of locomotion of Germanonautilus is reconstructed. Owing to the wide whorl cross section and the high whorl expansion rate, drag of the conchs was high, the aperture was oriented at an oblique angle which made Germanonautilus a rather slow horizontal swimmer. Because of their large sizes and widths, conchs of Germanonautilus were often deposited on their broad venters, forming elevated “benthic islands” (secondary hardgrounds). A broad range of animals (fish, decapods, ophiurans, crinoids, brachiopods, bryozoans, bivalves, Spirorbis, foraminiferans) lived in and on these comparatively large secondary hardgrounds.
A previously unreported large skull of the mastodonsaurid temnospondyl Xenotosuchus africanus, from the Cynognathus Assemblage Zone (Early to Middle Triassic) of the Beaufort Group, Karoo Basin of South Africa, is described. The species is plesiomorphic in most aspects of its cranial anatomy, and only one autapomorphy is tentatively identified herein, namely contact between the left and right exoccipitals posterior to the parasphenoid. The new specimen permits recognition of a growth series for X. africanus from sub−adult (all previously known specimens) to fully adult (new specimen) stages. Ontogenetic changes associated with this include changes in skull proportions, and changes in the morphology of the lacrimal flexure, frontal, postfrontal, cultriform process and body of the parasphenoid, and the transvomerine tooth row.
New records of nautiloid beak elements conventionally classified as 'Rhyncholithes hirundo (Biguet, 1819)' and 'Conchorhynchus avirostris (von Schlotheim, 1820)' with carbonised (originally chitinous) three-dimensionally preserved appendages from the Upper Muschelkalk (Middle Triassic) of northern Württemberg (Southwest Germany) enable restoration of the complete beak of Germanonautilus. In three specimens, the lower mandible is embedded within the living chamber of Germanonautilus conchs, Beak elements of Germanonautilus differ from those of Recent Nautilus in the more elongate appendages of the fossil lower mandibles and the weaker sculpture on the originally chitinous parts. Furthermore, the dorsal sculpture of the fossil conchorhynchs consists of ridges rather than denticles and the ventral sculpture of the fossil rhyncholiths displays ridges in places where the Recent rhyncholiths have a smooth surface. Additionally, the fossil beak elements attained a larger size than their Recent counterparts, During transport of 'Rhyncholithes hirundo', the light chitinous parts served as a sail and the heavier conchorhynch anchored in the sediment causing alignment. In contrast to the irregularly embedded isolated rhyncholiths, the conchorhynchs usually settled with their ventral side up. From the study of 407 fossil nautiloid beak-elements, a significant variability of the hard parts is evident. Consequently, the assignment of specific morphologies to the species of Germanonautilus is impossible.
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