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Crinoid ancestry without blastozoans

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At present, a debate in the paleontologic literature focuses on whether or not the immediate ancestry of the Crinoidea lies in an unidentified member of the Blastozoa, which includes eocrinoids and an assemblage known variously as the “cystoids”. Those proposing to derive crinoids from within the blastozoans have recently argued for homologies in the construction of the oral region of certain derived taxa from both groups. An opposing viewpoint, outlined here, finds evidence that aside from plesiomorphies, proposed similarities are superficial and homoplastic. We suggest these superficialities represent convergent adaptive strategies. Earliest crinoids express ambulacral traits unlike any blastozoan but that are expressed in the only other pentaradial echinoderms with a known record early enough to be considered in the context of crinoid origins, edrioasteroids and edrioasteroid-like stem echinoderms.
Loose elements of the roveacrinid Saccocoma from the Tithonian red Rogoża Coquina, Rogoźnik, Pieniny Klippen Belt, Poland, are used to test the contradictory opinions on the mode of life of Saccocoma. The investigated elements belong to three morphological groups, which represent at least two separate species: S. tenella, S. vernioryi, and a third form, whose brachials resemble those of S. vernioryi but are equipped with wings of different shape. The geometry of brachials’ articular surfaces reveals that the arms of Saccocoma were relatively inflexible in their proximal part and left the cup at an angle of no more than 45, then spread gradually to the sides. There is no evidence that the wings were permanently oriented in either horizontal or vertical position, as proposed by two different benthic life−style hypotheses. The first secundibrachial was probably more similar to the first primibrachial than to the third secundibrachial, in contrast to the traditional assumption. The winged parts of the arms were too close to the cup and presumably too stiff to propel the animal in the water efficiently. Swimming was probably achieved by movements of the distal, finely branched parts of the arms. The nonhorizontal attitude of the winged parts of the arms is also not entirely consistent with the assumption that they functioned as a parachute. Moreover, the wings added some weight and thus increased the energy costs associated with swimming. The hydrodynamic benefits balancing these extra costs are not entirely clear, but it seems probable that the wings reduced the sinking rate of the animal not by increasing the pressure drag, as suggested by the parachute−analogy, but by increasing the surface drag (friction drag), which also harmonize with the presence of spines, reticulate sculpture and conspicuous vacuolar ornamentation in some species of Saccocoma.
Disarticulated crinoid columnals and pluricolumnals from the Famennian of the Holy Cross area were analysed. Sixteen crinoid taxa were distinguished, only one of which is based on stems attributed to a calyx−based genus, and the others are classified within artificial supraspecific units. Two of these are new: Schyschcatocrinus levis sp. nov. and Cosmocrinus polonicussp. nov. The described crinoid fauna shows distinct extinction−recovery temporal pattern: the Frasnian–Famennian crisis affected 50% of stem−based families and 70% of late Frasnian stem−based genera. The succession of crinoid faunas represented by three faunal intervals has been identified and correlated to standard conodont zones: FIa, Palmatolepis triangularis Zone (relic “Frasnian” crinoid assemblage Schyschcatocrinus delicatus–Calleocrinus kielcensis), FIb, Pa. crepida to Pa. marginifera zones (crinoid assemblage Calleocrinus kielcensis–Schyschcatocrinus levis) and FIc, Pa. trachytera to S. praesulcata zones (crinoid assemblage Cosmocrinus polonicus–Acbastaucrinus affectatus). The succession was controlled mostly by eustatic factors.
The lecanocrinid Ammonicrinus(Flexibilia) is newly interpreted based on new material from the Middle Devonian of the Rhenish Massif (Eifel and Bergisches Land, Germany). The species have echinoid−like tubercles on the attachment and on the column, which bear articulated spines. The intraspecific variability of the column is discussed for three facies−controlled morphotypes, herein classified as standard “exposed−” or “encased roller−type” and the rare “settler−type”. New specimens have floating transitions between different plate sculpturing and between those individuals with none or one to several columnals with herein termed “lateral columnal enclosure extensions” on the proximal−most, barrel−like dististele and the following mesistele, which is solely distinguished by these extensions. Based on this interpretation, Ammonicrinus kongieli is evaluated as a subjective junior synonym of Ammonicrinus sulcatus. The latter species was first recognised from the Eifel (Germany). “Ammonicrinus wachtbergensis”, from the upper Eifelian of the Eifel, is declared a subjective junior synonym of Ammonicrinus doliiformis. The first nearly complete specimen of Ammonicrinus kerdreoletensis is described from the lower Eifelian of Vireux−Molhain (southern Ardennes, France). Two new species are described: Ammonicrinus jankei sp. nov. and Ammonicrinus leunisseni sp. nov. A functional morphologic trend in perfecting the crown encasement by continuous modification of the lateral columnal enclosure extensions of the mesistele from the Eifelian to the Givetian, indicates a vagile benthic “predator”−driven evolution of ammonicrinids in the Eifel area. Several ammonicrinid species are herein defined as spined soft−bottom dwellers, feeding in low−intensity current water, possibly through a self−produced water flow. The first known postmortem encrusting epizoans on ammonicrinid endoskeletons are reported.
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Crinoids from the Silurian of Western Estonia

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The Silurian crinoids of Estonia are re−evaluated based on new collections and museum holdings. Nineteen species−level crinoid taxa are now recognized. All crinoid names applied to Estonian Silurian crinoids during the middle 19th century are disregarded. Especially significant is the fauna reported herein from the Pridoli because coeval crinoids are very poorly known from the Baltic region and elsewhere. One new genus and four new species are described from Estonia, namely Calceocrinus balticensis sp. nov., Desmidocrinus laevigatus sp. nov., Eucalyptocrinites tumidus sp. nov., and Saaremaacrinus estoniensis gen. et sp. nov.
Bulk sampling of a number of different marine and marginal marine lithofacies in the British Bathonian has allowed us to assess the palaeoenvironmental distribution of crinoids for the first time. Although remains are largely fragmentary, many species have been identified by comparison with articulated specimens from elsewhere, whilst the large and unbiased sample sizes allowed assessment of relative proportions of different taxa. Results indicate that distribution of crinoids well corresponds to particular facies. Ossicles of Chariocrinus and Balanocrinus dominate in deeper−water and lower−energy facies, with the former extending further into shallower−water facies than the latter. Isocrinus dominates in shallower water carbonate facies, accompanied by rarer comatulids, and was also present in the more marine parts of lagoons. Pentacrinitesremains are abundant in very high−energy oolite shoal lithofacies. The presence of millericrinids within one, partly allochthonous lithofacies suggests the presence of an otherwise unknown hard substrate from which they have been transported. These results are compared to crinoid assemblages from other Mesozoic localities, and it is evident that the same morphological adaptations are present within crinoids from similar lithofacies throughout the Jurassic and Early Cretaceous.
Calcite isocrinid ossicles from the Middle Jurassic (Bathonian) clays in Gnaszyn (central Poland) show perfectly preserved micro− and nanostructural details typical of diagenetically unaltered echinoderm skeleton. Stereom pores are filled with ferroan calcite cements that sealed off the skeleton from diagenetic fluids and prevented structural and geochemical alteration. In contrast with high−Mg calcite skeleton of modern, tropical echinoderms, the fossil crinoid ossicles from Gnaszyn contain only 5.0–5.3 mole% of MgCO₃. This low Mg content can be a result of either a low temperature environment (ca. 10℃) and/or low Mg/Ca seawater ratio. Both conditions have been proposed for the Middle Jurassic marine environment. Occurrence of Mg−enriched central region of stereom bars of Jurassic columnal ossicle of Chariocrinus andreae is consistent with the concept of magnesium ions involvement in earliest growth phases of calcium carbonate biominerals.
Crinoid genus Ammonicrinus is represented by two species, A. sulcatus Kongiel and A. kongieli sp.n., in the Grzegorzowice - Skały and Świętomarz - Śniadka profiles of the Middle Devonian in the Holy Cross Mts. The presence of barrel-like columnals of a limited mobility in distal part of stems and the development of wide tuberculated external cover indicate that both species represent sessile benthos. In A. sulcatus stem was connected with crown through modified columnalium. The crown was equipped with short arms which could take food in a space limited to the interior of coiled part of the stem. The food was supplied by current parallel to the bottom. The structure of crown and stem suggests that all Ammonicrinus species represent the same evolutionary stage.
Early–Middle Frasnian ostracods and crinoids from Wietrznia in the Northern Kielce subregion of the Holy Cross area were analyzed. Twenty three ostracod species assigned to thirteen named genera, as well as eighteen crinoid species including the representatives of fifteen stem−based taxa were distinguished. For most of the species open nomenclature is applied. The composition of ostracod assemblage changes from moderately diverse in the lower part of the Palmatolepis transitans Zone to poorly diverse in its higher part. Lack of ostracods in the uppermost part of the Pa. transitans Zone and in the Palmatolepis punctata Zone is noted. The crinoid distribution pattern comprises the interval of relatively high diversity, interrupted in the uppermost part of the Pa. transitans Zone, and the interval of temporary recovery in the lower Pa. punctata Zone. Such distribution patterns point to deterioration of environmental conditions across the Early–Middle Frasnian transition, coinciding with a large−scale C−isotopic perturbation superimposed on intermittent, two−step eustatic sea level rise. On the other hand, impoverished, surviving crinoid faunas and absence of ostracods in the Pa. punctata Zone indicate the overall long−term deterioration of life conditions through the major C−isotope anomaly time span. However, this may also result from synsedimentary tectonic pulses, causing block movements and large−scale resedimentation phenomena on the northern slope of the Dyminy Reef during the basal Middle Frasnian sea level rise.
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