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n the goniatite family Prionoceratidae, the transition from Mimimitoceras to Balvia provides an example of rapid size decrease resulting from progenesis. In the Prionoceras-Mimimitoceras stock the adult conch continued to be of rather uniform shape and size (about 60 mm) and species diversification was expressed mostly in changing juvenile morphology. In the Balvia branch, which had developed in the Wocklumeria Stufe, the adult size diminished strongly (not more than 16 mm). Progenetic Balvia displays conch morphology of ancestral Mimimitoceras juveniles, with distinct ornamentation types that were added terminally.
Evolutionary lineages within the Carboniferous ammonoid superfamily Goniatitaceae can be recognized using cladistic and stratophenetic analyses, showing that both approaches lead to coinciding results. In the late Viséan and Namurian A, ammonoid provinces can be defined by the distribution of lineages within the goniatite superfamily Goniatitaceae. The first province corresponds to the Subvariscan Realm (where the superfamily became extinct near the Viséan-Namurian boundary), and the second embraces the majority of the occurrences, e.g. the south urals, central Asia, and North America (where the superfamily with different independent lieages continued up into the late Namurian A). In the Viséan, the superfamily was, in two short epochs, globally distributed with major transgressions, which probably led to migration events. The first is at the end of the late Viséan A (G. fimbriatus and G. spirifer Zones, when the genus Goniatites had a world-wide distribution with various species), and the second at the beginning of the late Viséan C (L. poststriatum Zone, when Lusitanoceras is globally distributed).
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The origin of ammonoid locomotion

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Evolution of the coiled ammonoid conch from the uncoiled bactritid conch was probably coupled with changes in manoeuvrability and swimming velocity. The gradual transformation of uncoiled to coiled ammonoid conchs has essential functional consequences. The radical change in conch geometry during phylogeny but also in ontogeny of early ammonoids implies a shift of the aperture from an original roughly downward, via a downward oblique and an upward oblique to an upward orientation, presuming a neutrally buoyant condition of the ammonoid animal. Similar trends were reconstructed for the three main ammonoid lineages in the Middle Devonian, the agoniatitid, the anarcestid, and the tornoceratid lineages. This allowed an increase in manoeuvrability and in the maximum horizontal swimming speed.
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The relatively rich assemblages of shark teeth from pelagic limestone (Mississippian, late Viséan, late Asbian-middle Brigantian) of three northern European regions: the Rhenish Mountains (Westenfeld Quarry, Germany), the Holy Cross Mountains (Todowa Grząba at the edge of Ostrówka Quarry, Poland), and Derbyshire (Cawdor Quarry, Matlock, England, UK) display certain similarities, with the absolute predominance of the teeth of Falcatidae (small Symmoriiformes) and the constant presence of Thrinacodus spp. The largest and most diverse assemblage from Todowa Grząba contains at least three species of a falcatid Denaea, a xenacanthimorph Bransonella nebraskensis, a newly described phoebodontid Thrinacodus dziki sp. nov., a few ctenacanthiform and euselachian teeth, and two abraded euchondrocephalan dental elements. Anachronistidae, common in the most of late Viséan pelagic faunas, are absent from Todowa Grząba and Westenfeld. The material under study differs from the shallow-water chondrichthyan fauna, hitherto described from the Mississippian carbonate platform facies, by its taxonomic content (particularly almost total absence of Euchondro-cephali), generally lower diversity, and higher frequency of small teeth.
Early late Viséan ammonoid assemblages in Morocco are composed of diverse and well-preserved specimens. The material was found in a plain in the Tafilalt (eastern Anti-Atlas). Here, we describe mass-occurrences of juvenile specimens, in which subadult and adult specimens occur in low numbers. The juveniles of some species display a conch morphology that differs fundamentally from the adult stages. Accordingly, we emend the species diagnoses of Goniatites lazarus as well as Calygirtyoceras darkaouaense, introduce the species Entogonites bucheri sp. nov., and discuss possible ecological implications of the morphologic changes throughout ontogeny. In particular, we compare the changes in conch morphology through ontogeny in the light of Pareto Optimiality according to which the morphology of organisms would fill a polygon or polyhedron in morphospace. Data points in one of the vorteces of the polyhedron indicate optimisation for the corresponding task. Although shape is not a proof of function, it appears plausible that juvenile conchs were selected rather for compactness while adult conchs were positively selected for conchs with improved hydrodynamic properties. This appears plausible because at small conch diameters, swimming movements will not suffice for effective translocation and a planktonic mode of life is likely.
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Raised shell projections on the inner shell walls that form pits on the internal moulds of Devonian ammonoids have been known for several decades. New specimens from Morocco reveal novel details of these structures; most, if not all, of which consist of a capsule of ammonoid shell that covers tiny tubes attached to the outer (= lateral or ventral) shell wall from the inside. In accordance with comparable Recent occurrences of similar structures in molluscs, we use the term “pearls” for these structures and the pits they form on the internal moulds. The nature of these encapsulated tubes is described and discussed. Because of the presence of these tubes inside the pearls, pearl arrangement, and their similarity to Recent mollusc occurrences, the tubes are interpreted as traces of parasitoses. The pearls and pits were grouped into five types based on differences in morphology, size, and arrangement. Then, having used these traits to perform a simple cladistic analysis, the resulting cladogram was compared to the phylogeny of ammonoids. Based on this comparison, it appears likely that the parasites underwent a co−evolution with the ammonoids, which lasted 10 to 15 Ma. Patterns of evolutionary events include co−speciation, “drowning on arrival” (end of parasite lineage near base of a new host clade), and “missing the boat” (parasite lineage does not adapt to a new host clade, thus not evolving a new parasite clade). Because of the lack of fossilised soft tissue, only speculations can be made about the systematic affiliation of the parasites, their life−cycle, infection strategy, and ecological framework. Some co−occurring bivalves also have pits reminiscent to structures caused by trematodes in Recent forms. Based on the available information, the tubes are interpreted as artefacts of trematode infestations, which, if correct, would extend the fossil record of parasitic trematodes into the Early Devonian.
An early Late Tournaisian (Early Carboniferous/Mississippian) ammonoid fauna is described from the Tafilalt of south−eastern Morocco. Twelve genera, four of which are new, and eleven new species are represented: Becanites africanus sp. nov., Triimitoceras epiwocklumeriforme gen. et sp. nov., Irinoceras minutum sp. nov., Muensteroceras quadriconstrictum sp. nov., Eurites bouhamedensis sp. nov., Ouaoufilalites ouaoufilalensis gen. et sp. nov., Helicocyclus fuscus sp. nov., Pericyclus mercatorius sp. nov., Orthocyclus(?) sp., Bouhamedites enigmaticus gen. et sp. nov., Winchelloceras antiatlanteum sp. nov., and Progoniatites maghribensis gen. et sp. nov. Palaeogeographic analysis of Late Tournaisian ammonoid assemblages shows strong endemism at the species−level, but genera and families had a nearly global distribution in the equatorial seas. The new fauna contains the stratigraphically oldest known representatives of the important Carboniferous goniatite families Girtyoceratidae and Goniatitidae.
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