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A Triassic spider from Italy

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A new fossil spider from the Triassic (Norian) Dolomia di Forni Formation of Friuli, Italy, is described as Friularachne rigoi gen. et sp. nov. This find brings the number of known Triassic spider species to four. The specimen is an adult male, and consideration of various features, including enlarged, porrect chelicerae, subequal leg length, and presence of a dorsal scutum, point to its identity as a possible member of the mygalomorph superfamily Atypoidea. If correct, this would extend the geological record of the superfamily some 98–115 Ma from the late Early Cretaceous (?Albian, c. 100–112 Ma) to the late middle–early late Norian (c. 210–215 Ma).
We report cyrtocrinid (Crinoidea) ossicles from the Rhaetian (Late Triassic) of the Tatra Mountains (southern Poland). The columnals are high, the facets are covered with thick crenulae and the latera are concave. Such features of symplectial articulation and latera distinguish them from the columnals of other Triassic crinoids (i.e., millericrinids and encrinids) and therefore we consider they belong to Cyrtocrinida. The oldest representatives of cyrtocrinids were known from the Early Jurassic, therefore the presented material constitutes the oldest world record of these crinoids to date. We speculate that perturbations related to the global mid−Carnian extinction combined with predation intensity observed in the Middle– Late Triassic have been involved in early origin of Cyrtocrinida.
Triassic corals with septa that branch repeatedly and centripetally are here assigned to a new genus Furcophyllia. Septa of F. septafindens (Volz, 1896), re−described from the Italian Dolomites, are composed of 3–10 blades (“septal brooms”). Distances between adjacent septa and their branches are equal, and the thickness of all blades is approximately the same throughout ontogeny. However, none of the septal brooms show the same branching pattern. Proposed herein is a simple computer model that reproduces septal pattern, similar to that of Furcophyllia, based on a minimal set of rules: (i) uniform coverage of intra−calicular space; (ii) regular bifurcations following some probability; (iii) keeping some minimal distance between septal branches. The elaborate septal pattern of Furcophyllia suggests a distinct organization of the polyp’s soft tissue, especially mesenteries whose appearance in modern corals is associated with insertion of sclerosepta. Hypothesis 1 suggests that mesenterial pairs flanked only “septal brooms” and that septal branches functionally corresponded with septal microarchitecture. Hypothesis 2 suggests that mesenterial pairs developed between all septal branches that functionally correspond with conventional septa. Delicate menianae, which developed on Furcophyllia septal faces (and many other Triassic corals) resemble similar septal microarchitecture of the Recent agariciid Leptoseris fragilis and may be closely related to the suspension feeding strategy of this coral. The furcate septal arrangement in Furcophyllia is unique among Triassic corals, and generally, among Mesozoic and Cenozoic corals. The only analogous corals are Cretaceous aulastraeoporids (e.g., Preverastrea, Paronastraea), Trochoidomeandra, and some Jurassic rhipidogyrids having secondary (apophysal) septal branches. In some Recent caryophylliids (Trochocyathus rhombocolumna, Phacelocyathus flos) primary septa may also split dichotomously and centripetally.
A histological analysis of the dentition of the small procolophonid parareptile Soturnia caliodon reveals detailed information concerning tooth implantation and replacement for this taxon. The presence of acrodont tooth implantation is verified, which contradicts current models for procolophonid dentition. A heterogeneous enamel layer, that reaches large thickness on the cusps, and a broad secondary dentine are also recorded. These structures provide a very stable occlusal morphology that extends the useful life of the teeth. During the process of replacement, old teeth were not pushed out but reabsorbed. The evidence indicates that Soturnia caliodon had a very low rate of tooth replacement which constitutes a valuable adaptation for its high−fibre herbivorous niche.
It is generally accepted that during the Triassic the composition of tetrapod faunas underwent a series of fundamental transformations, mainly as a result of diversification of archosaurs and decline of therapsids (Benton 1994, 2004, 2006). The last herbivorous basal synapsids, dicynodonts, disappeared from the record in the early Norian of the Americas, about 220 Ma (Langer et al. 2007), being unknown from the Late Triassic of Europe. Here, we report a partially articulated skeleton and isolated bones of a giant rhino−size dicynodont in the Upper Triassic fluvial sediments at Lisowice (Lipie Śląskie clay−pit) in southern Poland. Paleobotanical data indicate an early Rhaetian age for the fauna (Dzik et al. 2008; Niedźwiedzki and Sulej 2008). The dicynodont bones are associated with bones of carnivorous dinosaurs, pterosaurs, as well as capitosaur and plagiosaur amphibians. Dicynodonts were represented in the Germanic Basin throughout the Late Triassic, as proven by findings of smaller dicynodonts in older deposits in the same area, associated there with temnospondyl amphibians. It appears, thus, that the fossil record of tetrapod succession in the Late Triassic was strongly controlled by ecological factors and biased by uneven representation of particular environments. The Lisowice assemblage proves that faunas dominated by dicynodonts did not entirely disappear at least until the end of the Triassic.
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.
A recent contribution published in this journal (Dias−daSilva and Ilha 2009) reported a dermal skull fragment indicating the presence of a putative plagiosauroid temnospondyl in the Lower Triassic Sanga do Cabral Formation of the Paraná Basin, Southern Brazil. The taxonomic assignation of this specimen was necessarily tentative as it was based on circumstantial evidence, specifically the presence of a dense pustular ornamentation over four partial dermal skull bones, consideration of the described taxa known to bear such ornamentation, and the stratigraphic and paleobiogeographic range of such taxa. Therefore, Diasda−Silva and Ilha (2009) could not be totally certain about the plagiosauroid affinities of the new specimen and ascribed it to ?Plagiosauridae. It was particularly difficult to make a precise osteological identification of the specimen and six alternative osteological interpretations were made in comparison to both Gerrothorax and Peltobatrachus (see Dias−da−Silva and Ilha 2009: fig. 2). In spite of the poor taxonomic resolution, the new specimen raised interesting questions regarding the presence of plagiosauroid stereospondyls in western Gondwana, as well as their evolutionary patterns, biostratigraphic and paleobiogeographic implications. After Dias−da−Silva and Ilha's (2009) contribution was published, new data from Damiani et al. (2009) raised the possibility of narrowing down the taxonomic identity of the plagiosauroid from Brazil. Accordingly, this brief report provides a more precise taxonomic assignation for this material.
The Spathian (late Early Triassic) Virgin Formation of south−western Utah (USA) yields a comparatively diverse benthic fauna that flourished ~2 Ma after the end−Permian mass extinction. In this study, we present quantitative palaeoecological data, which are analysed in the context of depositional environments. This integrated approach helps to discriminate between effects of the end−Permian mass extinction event and local environmental factors on alpha diversity and ecological structure of the Virgin Fauna. Shallow subtidal environments yield the highest species richness and lowest dominance val− ues as recorded in two benthic associations, the Eumorphotis ericiusAssociation and the Protogusarella smithi Association, both ofwhich contain 20 benthic species (bivalves, gastropods, brachiopods, echinoderms, and porifers). Tidal inlet deposits yield a low diverse fauna (Piarorhynchella triassica Association) with a very high dominance of filter feeders adapted to high energy conditions.Another comparably low diverse fauna is recorded by the Bakevellia exporrecta Association, which occurs in deposits of the offshore transition zone,most likely reflecting unconsolidated substrates. A single sample contain− ing five bivalve species (Bakevellia costata Assemblage) is recorded from a marginal−marine setting. The Virgin fauna yields a bulk diversity of 30 benthic species (22 genera) of body fossils and 14 ichnogenera and, thus, represents the most di− verse marine bottom fauna known so far from the Early Triassic. Our results suggest that oceanographic conditions during the early Spathian enabled ecosystems to rediversify without major abiotic limitations. However, taxonomical differentia− tion between habitats was still low, indicating a time lag between increasing within−habitat diversity (alpha diversity) and the onset of taxonomical differentiation between habitats (beta diversity). We suggest that taxonomical habitat differentia− tion after mass extinction events starts only when within−habitat competition exceeds a certain threshold, which was not yet reached in the Spathian of the investigated area. This interpretation is an alternative to previous suggestions that the preva− lence of generalistic taxa in the aftermath of mass extinction events reflects protracted environmental stress. The onset of in− creasing beta diversity is a potential criterion for distinguishing two major recovery phases, the first ending with habitat satu− ration and the second ending with the completion of ecosystem differentiation.
The first osteohistological study focused exclusively on rhynchosaurs (non-archosauriform archosauromorphs), based on the hyperodapedontines Teyumbaita sulcognathus and Hyperodapedon sp., from the Upper Triassic of Southern Brazil, indicates a relatively rapid growth rate in early ontogeny shown by the fibrolamellar complex, with a change to slow intermittent growth during late ontogeny represented by parallel-fibred bone with several growth marks. Contrary to previous studies, which described a typical non-archosaur reptilian bone tissue pattern for rhynchosaurs, with growth marks extending across the entire cortex, we demonstrate that, in both studied taxa, the initial growth rate was faster in comparison to the later. This suggests that the ability of rapid growth at high rates was already present in basal non-ar-chosauriform archosauromorphs.
Postcranial skeletal pneumaticity (PSP) is present in a range of basal sauropodomorphs spanning the basal sauropodomorph–sauropod transition. We describe the PSP of five taxa, Plateosaurus engelhardti, Eucnemesaurus fortis, Aardonyx celestae, Antetonitrus ingenipes, and an unnamed basal sauropod from Spion Kop, South Africa (hereafter referred to as the Spion Kop sauropod). The PSP of Plateosaurus is apparently sporadic in its occurrence and has only been observed in very few specimens, in which it is of very limited extent, affecting only the posterior cervical vertebrae and possibly the mid dorsals in one specimen. The PSP of Eucnemesaurus, Aardonyx, Antetonitrus, and the Spion Kop sauropod consists of subfossae (fossa−within−fossa structures) that excavate the vertices of the posterior infradiapophyseal fossae of the posterior dorsal vertebrae. These subfossae range from simple shallow depressions (Eucnemesaurus) to deep, steepsided, internally subdivided and asymmetrically developed chambers (Antetonitrus). The middle and anterior dorsal vertebrae of these taxa lack PSP, demonstrating that abdominal air sacs were the source of the invasive diverticula. The presence of pneumatic features within the infradiapophyseal fossae suggest that the homologous fossae of more basal saurischians and dinosauriforms were receptacles that housed pneumatic diverticula. We suggest that it is probable that rigid non−compliant lungs ventilated by compliant posterior air sacs evolved prior to the origination of Dinosauria.
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.
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New species of megaspores from the Trias of Poland

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Thirty new species and two new genera (Otynisporites, Bothriotriletes) of megaspores from the Trias of Polish Lowlands have been described. Most of them are from the Lower Keuper (Lettenkohle) and the Upper Buntsandstein. The megaspores from the Lower Buntsandstein have been described.
A new generic name, Cycliphyllia, is here proposed as a replacement name for Cyclophyllia Roniewiecz, 1989 (type species: Thecosmilia cyclica Schaefer et Senowbari-Daryan, 1978, Upper Triassic). The latter is a junior homonyme of Cyclophyllia Milne-Edwards et Haime, 1848 (type species: Cyclolites cristata Lamarck, 1801, Cretaceous), an invalid name, which is a junior synonyme of Aspidiscus Koenig, 1825 (Milne-Edwards 1857: t. 2, p. 386). This regrettable error has been noticed thanks to the List of generic names by Wells (1986). As a consequence of the above change, the orthography of the family name Cyclophylliidae Roniewicz, 1989 is here corrected into Cycliphylliidae.
The Middle Triassic ammonoid genus Ceratites diversified spectacularly within the Germanic Muschelkalk Basin during the Anisian/Ladian (244–232 Mya). Previous studies have interpreted this diversification as a sequence of rapid, endemic radiations from a few immigrant taxa. Here we investigate the possibility that geological and sampling biases, rather than ecological and evolutionary processes, are responsible for this pattern. A new specimen−based dataset of Ceratites species−richness and abundance was assembled. This dataset was combined with 1:200 000 geological maps in a geodatabase to facilitate geospatial analyses. One set of analyses compared species richness per geological map with the number of occurrences and localities per map. Per−map change in the amount of rock available to sample for fossils was also included as a variable. Of these three variables, number of occurrences is the most strongly correlated with richness. Variation in the amount of rock is not a strong determinant of species−richness. However, rarefaction of basin−wide species/abundance data demonstrates that differences in species−richness through time are not attributable to sample size differences. The average percent similarity among sites remained close to 50% throughout the Upper Muschelkalk. The rank abundance distribution (RAD) of species from the first interval of the Upper Muschelkalk is consistent with colonization of a disturbed environment, while the other two intervals have RADs consistent with more stable ecosystems. These results indicate that genuine ecological and evolutionary events are partly responsible for the observed differences in richness and abundance. Although changes in the RADs through time support changes in the ammonoid assemblage structure, the processes underlying increasing richness and change in RADs cannot be explained by increasing geographic distinctiveness or isolation among the ammonoid assemblages present at different localities.
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.
The Temnospondyli are a large and diverse group of stemtetrapods (sensu Laurin and Reisz 1997) known from the Early Carboniferous to the Early Cretaceous; their remains have been found on all continents, from Greenland to Antarctica. The Metoposauridae are a short−ranging temnospondyl group recorded only from the Late Triassic; they are characterized by an anterior position of the orbits, a broad parasphenoid and a large quadrate foramen. However, unclear is the taxonomic value of the external location of tusks on the dentary. Our analysis of tooth rows in Metoposaurus diagnosticus krasiejowensis from the Triassic (Late Carnian) of Krasiejów (Silesia, SW Poland) shows that the external location of tusks on the dentary is not a synapomorphy for Metoposauridae but a character of great intraspecific variability. Variability of the arrangement of the internal tooth row on the upper jaw has also been observed.
Re−investigation of the skull roof in Metoposaurus diagnosticus from the German Middle Keuper revealed that in contrast to previous opinions, the lacrimal bone in this species enters the orbital margin.The same pattern is demonstrated by the skulls of a newly discovered metoposaur from the Keuper of Krasiejów in Poland.The difference in the shape of the parietal between the population from Krasiejów and the type population of Metoposaurus diagnosticus enables the discrimination of two separate subspecies within Metoposaurus diagnosticus.For the specimens from the Late Carnian of Drawno Beds at Krasiejów, Poland and its lateral equivalents Lehrberg Beds at Stuttgart−Sonnenberg and Kieselsandstein at Fichtenberg, Germany, a new chronosubspecies Metoposaurus diagnosticus krasiejowensis is erected.The new subspecies differs from the older nominal subspecies M. diagnosticus diagnosticus in having a shorter and wider prepineal part of the parietal.If one accepts that the nominal subspecies is the ancestor of M. krasiejowensisthe change in the shape of the parietal would be a reversal of the trend towards elongating postorbital part of the skull observed in ancestors of the metoposaurids.It seems that the skull development in ontogeny changed after the anterior shift of the orbits occurred in the phylogenetic history of the metoposaurids.The difference in ornamentation of the interclavicle between European Metoposaurus and North American genera is corroborated by Polish material.
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