Recovery dynamics of benthic marine communities from the Lower Triassic Werfen Formation,northern Italy

The Lower Triassic Werfen Formation of northern Italy represents an important archive for Early Triassic ecosystems. Based on quantitative community analysis using species level identifications, we reconstruct the recovery of benthic ecosystems after the end‐Permian mass extinction throughout this unit. The analysis of benthic macrofossil communities shows that incipient recovery has taken place during the Griesbachian and the Griesbachian–Dienerian transition. A probable deterioration in environmental conditions is observed towards the end of the Dienerian. The Smithian part of the Werfen Formation is characterized by high siliciclastic input, which ceases around the Smithian‐Spathian boundary. The Spathian marks the definitive phase of recovery in the Werfen Formation. The comparison of this pattern with other palaoegeographical regions suggests that both the Griesbachian recovery and the Dienerian decline were of inter‐regional if not global extent, whereas the Smithian diversity low in the Werfen Formation is a local signal. In contrast to the recovery dynamics of ammonoids and conodonts, the Smithian–Spathian boundary interval was no caesura for benthic ecosystems. The Spathian recovery pulse is possibly also an inter‐regional event, at least in the palaeotropics. These results are in contrast with the previously proposed scenario of persistent hostile conditions during the Griesbachian time interval and highlight an initial recovery phase restricted to Griesbachian times. Instead, the apparently sluggish recovery of benthic ecosystems was at least partly shaped by set‐backs due to short‐term environmental perturbations during the Dienerian.     

Diversity and palaeoecology of a highly diverse Late Triassic marine biota from the Cassian Formation of north Italy

A sample of marine invertebrates from the Late Triassic Cassian Formation (north Italy) yielded one of the most diverse Early Mesozoic fossil assemblages ever reported (c. 170 species). The assemblage was found in basin clays, but was transported from nearby carbonate platforms as indicated by fragmentation, microbial encrustation and the presence of coated grains and ooids. Most of the specimens are small (< 1 cm) reflecting both, small adult sizes and size sorting during transport. Rarefaction analysis suggests that diversity of surface collection and bulk sampling is the same. However, rank abundance, species richness and taxonomic composition differ strongly according to sampling method. Low‐grade lithification of the sediments is the main reason that high diversity can be recognized, because it facilitates disaggregation and finding of small molluscs. Sample standardization shows that the studied assemblage is much more diverse than known Early Triassic assemblages. However, its diversity is similar to that of Anisian assemblages. This suggests that recovery from the end‐Permian mass‐extinction was quite advanced in the Middle Triassic and alpha‐diversity remained high until the Late Triassic. According to current models, Early Triassic and Anisian faunas match the niche overlap phase of recovery during which diversity is built up by increasing alpha‐diversity, whereas beta‐diversity rises slowly. Subsequently, habitat width of species contracts because of increasing competition, making beta‐diversity the principal drive of overall diversity increase. The diversity pattern of various Late Triassic Cassian associations meets the predictions for the transition from the niche overlap to the habitat contraction phase.: Triassic, Cassian Formation, palaeoecology, diversity, mollusc dominance.     

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.     

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).     

三叠纪胎生主龙型海洋爬行动物

现代爬行动物的生殖方式包括卵生、卵胎生和“胎生”三种类型.总体而言,卵生即母体在陆地上产出羊膜卵,并依靠外界环境加以孵化;卵胎生系指羊膜卵在母体内孵化,随即产出幼子,其间胚胎的营养来源和代谢废物均限于羊膜卵内;所谓“胎生”,是指某些类群的卵在母体内孵化后没有立刻产出,胚胎与母体建立了某种程度的营养乃至气体交换关系,此后再以幼体的形式娩出.卵胎生和“胎生”见于有鳞类.在化石中,由于无法识别胚胎与母体的营养联系等信息,通常只以“生产方式”加以定义,即除卵生以外,所有“含胚胎的化石”都统称为“胎生”,目前“胎生”已有的准确记录包括鱼龙类、鳍龙类、沧龙类和离龙类.近期报道的“怀孕恐头龙”显示原龙类可能也有此种生殖方式,但标本的保存状态显示不能排除其“同类相食”之可能.本文记述了产自云南罗平地区中三叠系内一独立保存的、完整的圆胚状化石,标本的大小、形态、姿态和埋藏环境显示这是一个处于孵化后期、因某种原因而流产的胚胎.头后骨骼显示该标本代表了一种全新类型的海生原龙类,但是由于在个体发育中很多性状特征会有较大变化,因此胚胎乃至幼体标本不宜作为正型标本而建立新属种.这也是又一个主龙型爬行动物胎生的确切证据;同时这也进一步反映了盘县—罗平动物群中原龙类的高度多样性.     

The Luoping biota: exceptional preservation, and new evidence on the Triassic recovery from end-Permian mass extinction

The timing and nature of biotic recovery from the devastating end-Permian mass extinction (252 Ma) are much debated. New studies in South China suggest that complex marine ecosystems did not become re-established until the middle–late Anisian (Middle Triassic), much later than had been proposed by some. The recently discovered exceptionally preserved Luoping biota from the Anisian Stage of the Middle Triassic, Yunnan Province and southwest China shows this final stage of community assembly on the continental shelf. The fossil assemblage is a mixture of marine animals, including abundant lightly sclerotized arthropods, associated with fishes, marine reptiles, bivalves, gastropods, belemnoids, ammonoids, echinoderms, brachiopods, conodonts and foraminifers, as well as plants and rare arthropods from nearby land. In some ways, the Luoping biota rebuilt the framework of the pre-extinction latest Permian marine ecosystem, but it differed too in profound ways. New trophic levels were introduced, most notably among top predators in the form of the diverse marine reptiles that had no evident analogues in the Late Permian. The Luoping biota is one of the most diverse Triassic marine fossil Lagerstätten in the world, providing a new and early window on recovery and radiation of Triassic marine ecosystems some 10 Myr after the end-Permian mass extinction.     

The role of the forelimb in prey capture in the Late Triassic reptile Megalancosaurus (Diapsida,Drepanosauromorpha)

The pectoral girdle and forelimb of the Late Triassic drepanosauromorph reptile Megalancosaurus are redescribed and their function reinterpreted. The whole skeleton of this diapsid is highly specialised for arboreal life, and also the peculiarities of the shoulder girdle and forelimb were interpreted as adaptations for a limb-based locomotion using gap-bridging to move from one support to another, as in chameleons. Re-examination of the pectoral girdle and forelimb revealed the presence of clavicles fused into a furcula-like structure, a saddle-shaped glenoid and a tight connection between the radius and ulna that strengthened the forearm but hindered pronation and supination movements at that joint. The new information plus a reconstruction of the pectoral and forelimb musculature suggests that the forelimb was also specialised for grasping and raking in addition to climbing and thus prey capture may have been an important function for the forelimb. The new functional interpretation fits well with the overall body architecture of Megalancosaurus’ skeleton, suggesting that this reptile was an ambush predator that may have assumed a stable tripodal position, secured by the hooked tail and hind limbs, freeing its forelimbs to catch prey by sudden extension of the arm and firm grasping with the pincer-like digits.     

Dasycladalean green algae from the Upper Triassic of Mt. Rotonda (Verbicaro Unit, Calabria-Lucania Border, Southern Italy)

Summary A rich and diverse dasycladalean algae association is described from the Upper Triassic succession of Mt. Rotonda (Calabria-Lucania border, Southern Italy). This association consists of:Neoteutloporella rajkae n.sp.,Griphoporella bechst?dti n.sp.,Physoporella zamparelliae n.sp.,Spinaporella andalusica Flügel & Flügel-Kahler, 1984,S.? granadaensis Flügel & Flügel-Kahler, 1984,Chinianella? sp.,Gyroporella sp.,Griphoporella? sp. andPhysoporella aff.leptotheca. Neoteutloporella rajkae n.sp. is characterised by an undulated calcareous skeleton with short acrophore primary laterals bearing a tuft of 4–6 elongate, segmented, trichophore secondary laterals. This species allows to extend back to the Upper Triassic the stratigraphic range of the genusNeoteutloporella, previously known only from Upper Jurassic levels. Griphoporella bechst?dti n.sp. has a cylindrical calcareous skeleton and primary laterals only, consisting of a thin proximal part followed by a swollen portion that pinches out distally and finally opens outward with a cup-like swelling. Physoporella zamparelliae n.sp. is characterised by a calcareous skeleton made by partly welded thin individual sheaths enclosing the laterals. The laterals are piriferous, vertically compressed, roughly triangular both in vertical and in verticillar section. In some specimens they end with a spine-like thin apophysis. This species confirms that the typical Middle Triassic genusPhysoporella survived up into the Norian. The dasycladalean algal association of the Norian of Mt. Rotonda shows some similarities with the algal association found in the Upper Triassic of the Betic Cordillera whereas it is markedly different from the rich association occurring in the Upper Triassic of Sicily and of the Northern Calcareous Alps. This pattern is coupled with a different composition of the platform margin communities: microbial/serpulids bioconstructions in the Upper Triassic of the Calabria-Lucania border and of Alpujarridevs. Dachstein-type reefs in Sicily and the Northern Calcareous Alps. This indicates that the palaeoceanographic and palaeogeographic conditions controlled both the development of the different platform margin and of the different algal assemblages.     

Pseudoplanktonic lifestyle of the Triassic crinoid Traumatocrinus from Southwest China

Colonies of Traumatocrinus (Echinodermata, Crinoidea) attached to driftwood from the lower Upper Triassic (Carnian) Xiaowa Formation of the Guanling area (Guizhou, Southwest China) document a pseudoplanktonic lifestyle for this specialized offshoot of the otherwise benthic Middle Triassic Encrinidae. After the end-Carnian decline of Traumatocrinus, its ecological niche was subsumed in Norian by genera derived from another benthic family (Holocrinidae) with convergent morphological modifications. After the Toarcian (Lower Jurassic), this niche disappeared, possibly due to the emergence of wood-boring bivalves.     

New dasycladalean algae from the Middle Norian (Upper Triassic) of northern Italy (Mt. Pramaggiore, Carnic Prealps)

In the Eastern Southern Alps of northern Italy (Carnic Prealps, Friuli region), the shallow-water carbonate platform deposits of the Dolomia Principale Fm. (Norian–Rhaetian, Upper Triassic) show best-preserved platform to basin facies transition. The palaeontological study of an algal-rich level recovered from the platform margin facies (Mt. Pramaggiore) has displayed a very interesting association of Dasycladales. Two new genera (Bystrickyella and Elliottporella) and four new species (Bystrickyella ottii, Elliottporella morelloae, Palaeodasycladus lorigae and Holosporella conradii) have been described. These new data suggest that the Norian represents a period of turnover in the evolutionary history of the green algae community. This stage, placed between two extinctions, end-Ladinian and end-Norian, is here interpreted as a re-organization period of the evolutionary schemes of Dasycladales. The new lineages originated in the Norian developed further and characterized the Early Jurassic scenery. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

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 .     

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.     

Dasycladalean algae distribution in ammonoid-bearing Middle Triassic platforms (Dolomites, Italy)

This paper gives an account of distribution of Dasycladales in ammonoid-bearing Middle Triassic carbonate platforms of the Dolomites. A ca. 600-m-thick section was investigated within the Latemar platform interior and Dasycladales from correlated ammonoid-bearing samples of the Marmolada platform were studied for comparison. At Latemar, the dense sampling allowed, with some confidence, the identification of the first occurrences of Diplopora nodosa and Gyroporella ladinica, and the last occurrences of D. annulatissima and D. comelicana. Their stratigraphic range was found to be in agreement with literature. Other first or last occurrences could not be reconciled with known distributions, and were considered as environmentally or palaeogeographically controlled. All four algal events occur within a narrow interval of two Illyrian (Upper Anisian) ammonoid subzones (avisianum and crassus subzones). This study demonstrates the biostratigraphic potential of Dasycladales in platform settings, and highlights the necessity of detailed stratigraphic studies to determine their distribution.     

An enigmatic marine reptile,Hispaniasaurus cranioelongatus (gen. et sp. nov.) with nothosauroid affinities from the Ladinian of the Iberian Range (Spain)

An incomplete skull of a marine reptile with an atypical elongation of the postorbital region is described. The find comes from the Muschelkalk facies (Cañete Formation) of the Villora section (Iberian Range, Cuenca Province, Spain), characterised by a shallow marine (intertidal) environment and dated as Ladinian in age. The small skull has a rectangular shape, lacking, as preserved, upper temporal openings and a parietal foramen. The upper temporal openings might be secondarily closed. However, the absence of a parietal foramen and squamosals in the preserved part and the incompleteness of the pterygoids make a posteriorly postponed location of the upper temporal openings also conceivable. Teeth are all broken off but alveolar spaces indicate large and massive maxillary dentition. Micro-CT-data revealed a highly vascularised inner structure of the dorsal skull elements, which might indicate special feeding adaptations. Adding the new find to an existing phylogenetic analysis of Triassic marine reptiles reveals eosauropterygian, especially nothosauroid, affinities. However, morphological differences to nothosauroids justify the erection of a new genus and species for this enigmatic marine reptile. Its atypical morphology, without any extinct or modern analogue, fits well with the continuously increasing diversity of Triassic marine reptiles, exhibiting various specialised feeding strategies urn:lsid:zoobank.org:act:6D75AEC7-A5C5-4844-B71A-8215AB099134     

Lignicolous marine fungi in the Straits of Messina,Italy

The occurrence and distribution of lignicolous marine fungi in the Straits of Messina has been studied. Using submerged panels of pine, beech and poplar, twenty fungal species were identified. The lignicolous mycoflora of the Messina Straits was not significantly different from that reported in the literature for other temperate marine coastal environments, Ascomycotina being frequent, while Basidiomycotina were rare.     

Lycophytes from the Middle Triassic (Anisian) locality Kühwiesenkopf (Monte Prà della Vacca) in the Dolomites (northern Italy)

Abstract: Generally, lycophytes are not well represented in Anisian floras. Nonetheless, four different genera, each with one species, have been distinguished in the Anisian flora from Kühwiesenkopf (Monte Prà della Vacca), Italy. They were well preserved and yielded leaf cuticles in all four taxa and in situ spores in two of them. Lycopia dezanchei gen. et sp. nov. is the most common form; it is characterized by a dichotomising creeping prostrate rhizome (representing the primary shoot axis) from which arise aerial axes, apically bifurcated and covered with bundles of long leaves. This taxon cannot be attributed to any of the lycophyte orders, but most resembles Lycopodiales. Two representatives of the Isoetales have been found, of which Isoetites brandneri sp. nov. is abundant. This species is characterized by a short stem, which is unusual for this genus. In situ immature microspores and megaspores were found in some specimens. Lepacyclotes bechstaedtii sp. nov. has a quadrilobe corm with fertile scales inserted in whorls or slightly helicoidally. Short sterile leaves arise distally from the corm in some specimens. Selaginellales are very rare in the flora; Selaginellites leonardii sp. nov. is represented by a strobilus that yielded both micro‐ and megaspores. Some sterile fragments have been found associated, but never in organic connection.