Sequence of post-moult exoskeleton hardening preserved in a trilobite mass moult assemblage from the Lower Ordovician Fezouata Konservat-Lagerstätte, Morocco

Euarthropods have a tough exoskeleton that provides crucial protection from predation and parasitism. However, this is restrictive to growth and must be periodically moulted. The moulting sequence is well-known from extant arthropods, consisting of: (i) the long inter-moult stage, in which no changes occur to the hardened exoskeleton; (ii) the pre-moult stage where the old exoskeleton is detached and the new one secreted; (iii) exuviation, when the old exoskeleton is moulted; and (iv) the post-moult stage during which the new exoskeleton starts as soft, thin, and partially compressed and gradually hardens to the robust exoskeleton of the inter-moult stage. Trilobite fossils typically consist of inter-moult carcasses or moulted exuviae, but specimens preserving the post-moult stage are rare. Here we describe nine specimens assigned to Symphysurus ebbestadi representing the first group of contemporaneous fossils collected that preserve all key stages of the moulting process in one taxon, including the post-moult stage. They were collected from a single lens in the Tremadocian part of the Fezouata Shale Formation, Morocco. Based on cephalic displacement and comparison to other trilobite moults, one specimen appears to represent a moulted exoskeleton. Four specimens are typical inter-moult carcasses. Four others are wrinkled and flattened, with thin exoskeletons compared to inter-moult specimens, and are considered post-moult individuals. These S. ebbestadi specimens illuminate the preservation and morphology of the post-moulting stage, characterised by strong anterior-posterior exoskeleton wrinkling, as well as overall body flattening and reduced visibility of thoracic articulations. Being found in the same lens, these specimens likely represent the first preserved in-the-act mass moulting event. The displayed sequence of moulting suggests the moulting process in trilobites was comparable to modern arthropods, and conserved within euarthropod evolutionary history

Phytoplankton dynamics from the Cambrian Explosion to the onset of the Great Ordovician Biodiversification Event: a review of Cambrian acritarch diversity

Most early Palaeozoic acritarchs are thought to represent a part of the marine phytoplankton and so constituted a significant element at the base of the marine trophic chain during the ‘Cambrian Explosion’ and the subsequent ‘Great Ordovician Biodiversification Event.’ Cambrian acritarch occurrences have been recorded in a great number of studies. In this paper, published data on Cambrian acritarchs are assembled in order to reconstruct taxonomic diversity trends that can be compared with the biodiversity of marine invertebrates. We compile a database and calculate various diversity indices at global and regional (i.e. Gondwana or Baltica) scales. The stratigraphic bins applied are at the level of the ten Cambrian stages, or of fourteen commonly used biozones in a somewhat higher resolved scheme. Our results show marked differences between palaeogeographical regions. They also indicate limitations of the data and a potential sampling bias, as the taxonomic diversity indices of species are significantly correlated with the number of studies per stratigraphic bin. The total and normalized diversities of genera are not affected in the same way. The normalized genus diversity curves show a slow but irregular rise over the course of the Cambrian. These also are the least biased. A radiation of species and to a lesser extent of genera in the ‘lower’ Cambrian Series 2 appears to mirror the ‘Cambrian Explosion’ of metazoans. This radiation, not evident on Gondwana, is followed by a prominent low in species diversity in the upper Series 3 and lower Furongian. Highest diversities are reached globally, and on both Baltica and Gondwana, in the uppermost Cambrian Stage 10, more precisely in the Peltura trilobite Zone, preceding a substantial phase of acritarch species extinction below and at the Cambrian/Ordovician boundary. Nearly all the genera present in Stage 10 survived into the Ordovician. The forms that emerged during the Cambrian therefore became the foundation for the more rapid radiation of acritarchs during the ‘Great Ordovician Biodiversification Event’

Reconstructing depositional rates and their effect on paleoenvironmental proxies : the case of the Lau Carbon Isotope Excursion in Gotland, Sweden

Variations in depositional rates affect the temporal depositional resolutions of proxies used for paleoenvironmental reconstructions; for example, condensation can make reconstructed environmental changes appear very abrupt. This is commonly addressed by transforming proxy data using age models, but this approach is limited to situations where numerical ages are available or can be reliably inferred by correlation. Here we propose a new solution, in which relative age models are constructed based on proxies for depositional rates. As a case study, we use the onset of the late Silurian Lau Carbon Isotope Excursion (LCIE) in Gotland, Sweden. The studied succession is a gradual record of shallowing upward in a tropical, neritic carbonate platform. As proxies for depositional rates we tested thorium concentration, carbonate content, and the concentration of pelagic palynomorphs. These three proxies were used to create relative age models using the previously published DAIME model. We applied these models to transform the delta C-13(carb) values as well as concentrations of selected redox-sensitive elements. The three relative age models yielded qualitatively similar results. In our case study, variations in depositional rates resulted in peaks of redox proxies appearing up to 76% higher when taken at face value, compared to when accounting for these rates. In the most extreme cases, our corrections resulted in a reversal in the stratigraphic trend of elemental concentrations. This approach can be applied and developed across depositional setting and types of paleoenvironmental proxies. It provides a flexible tool for developing quantitative models to improve our understanding of the stratigraphic record

Graptolites from Silurian (Llandovery Series) sedimentary deposits attributed to a forearc setting, Co To Formation, Co To archipelago, northeast Vietnam

Newly collected graptolites from the Co To Formation, Co To archipelago, NE Vietnam, comprise assemblages indicative of two biostratigraphical levels within the lower Silurian, Llandovery Series, Telychian Stage: the co-occurrence of Spirograptus turriculatus and Torquigraptus proteus? suggests an interval most likely within the upper part of the Spirograptus turriculatus Biozone or 'Monograptus' crispus Biozone, whilst Oktavites spirally and Monoclimacis cf. suhgeinitzi identify the Oktavites spiralis Biozone. The graptolites provide important biostratigraphical evidence for the age of the upper part of the lower Co To Formation, biostratigraphical ties between the NE Vietnamese succession of the Bac Bo Region and graptolite assemblages of the Long Dai Formation in the Viet-Lao Region of central Vietnam, and include the new species Monograptus hanutlus sp. nov. co-occurring with S. turriculatus, which is perhaps an ancestral form to the later Telychian species Monograptus drepanoformis. We also report the lirst chitinozoans, including Belonechitina, from the Co To Formation

The rise of pinnacle reefs : a step change in marine evolution triggered by perturbation of the global carbon cycle

The first appearance of pinnacle reef tracts, composed of hundreds to thousands of localized biogenic structures protruding tens to hundreds of meters above the surrounding mid-Silurian seafloor, represents a step change in the evolution of the marine biosphere. This change in seafloor morphology opened a host of new ecological niches that served as "evolutionary cradles" for organism diversification. However, the exact timing and driver's of this event remain poorly understood. These uncertainties remain, in large part, due to a paucity of index fossils in the reef facies, the difficulty of correlating between the offshore pinnacle reefs and more temporally well-constrained shallow marine fades, and cryptic unconformities that separate amalgamated reefs. Here we use delta C-13(carb) stratigraphy within a sequence stratigraphic framework to unravel these complex relationships and constrain the origination of Silurian pinnacle reef tracts in the North American midcontinent to near the Pt. celloni Superzone-Pt. am. amorphognathoides Zonal Group boundary of the mid-Telychian Stage. In addition, we identify a striking relationship between pulses of reef development and changes in global delta C-13(carb) values and sea level. Viewed through this new perspective, we correlate prolific periods of reef development with short-lived carbon isotope (delta C-13(carb)) excursions and eustatic sea level change that, ultimately, reflect perturbations to the global carbon cycle. From changes in the dominance of microbial reefs of the Cambrian to metazoan colonization of reefs in the Middle Ordovician, through the subsequent collapse of metazoan diversity with the Late Ordovician mass extinction, and the first appearance of early Silurian (Llandovery) pinnacle reef tracts and their proliferation during the late Silurian (Wenlock-Pridoli) and Devonian, major reef formation intervals increasingly coincide with delta C-13(carb) excursions. These patterns suggest that Paleozoic reef evolution was the product of environmental forcing by perturbations of the global carbon cycle

Complex response of dinoflagellate cyst distribution patterns to cooler early Oligocene oceans

Previous studies have made extensive use of dinoflagellate cysts to reconstruct past sea surface temperature (SST). Analysis of associations of dinoflagellate cysts using two new ocean datasets for the mid Eocene (Bartonian) and early Oligocene (Rupelian) reveals clear latitudinally constrained distributions for the Bartonian, but unexpected changes in their Rupelian distribution; a significant number of species with low and mid latitude northern hemisphere occurrences in the Bartonian extend their northward ranges in the Rupelian, including some forms characterised as ‘warm water’ by previous studies. This suggests either that dinoflagellates are faithfully tracking a complex oceanographic response to Rupelian cooling, or that dinoflagellate sensitivity/adaptability to a range of ecological variables means that at a global scale their distributions are not primarily controlled by sea surface temperature-variability. Previous use of dinoflagellate cysts for palaeoclimate work has relied on rather subjective and inconsistent identification of ‘warm’ and ‘cold’ water forms, rather than comprehensive analysis of community associations at the global-scale. It is clear from this study that a better understanding of the (palaeo-)ecology of dinoflagellates and their cysts is required. Rupelian dinoflagellate cyst distribution may reflect changes in a range of environmental variables linked to early Oligocene climate-cooling, for example changes in nutrient fluxes triggered by glacially-induced base-level fall; complex reorganisation of ocean current systems between the Bartonian and Rupelian, or muted changes to Rupelian summer SSTs in the northern hemisphere that have previously been reported. Many extant dinoflagellate species also exhibit relatively broad temperature tolerance. Moreover, they have potentially extensive cryptic diversity, and are able to produce dormant cysts during short-lived environmental deterioration, all of which may act to limit the value of undifferentiated dinoflagellate cyst assemblages for identifying climate signals

Gauging the impact of glacioeustasy on a mid-latitude early Silurian basin margin, mid Wales, UK

The early Silurian (Llandovery) Gondwanan South Polar ice sheet experienced episodes of ice retreat and re-advance. Marine base level curves constructed for the interval are widely assumed to provide a record of the associated glacioeustasy. In revealing a series of progradational sequences (progrades) bounded by flooding surfaces, recent work on the Type Llandovery succession in mid Wales (UK) has provided an opportunity to test this hypothesis. The grouping of these progrades into three composite sequences underpins the construction of both low order (small amplitude, high frequency) and high order (large amplitude, low frequency) base level movement curves. Revised biostratigraphical datasets for the type succession permit the accurate dating of base level events. The composite sequences record progradational acmes in the acinaces, lower convolutus and upper sedgwickii-halli graptolite biozones. A series of transgressions that postdate the Hirnantian glacial maximum culminated in an upper persculptus Biozone high-stand. Maximum flooding events also occurred during the revolutus and lower sedgwickii biozones, and the base of the early Telychian guerichi Biozone also marked the onset of a pronounced deepening. A review of 62 published datasets, including global and other regional base level curves, records of glacial activity, isotope data, patterns of facies and faunal flux and putative climate models, permits an evaluation of the origins of these local base level events. The concept of a Eustasy Index is introduced and shows that the impacts of global sea level movements can only be demonstrated within narrow ‘eustatic windows’ coincident with times of ice sheet collapse. At other times, the geometry of Llandovery area progrades reflects their accumulation across a faulted basin margin where, during periods of slow ice sheet advance, epeirogenic processes outstripped sea level movements as the dominant forcing factors. Increased levels of Telychian subsidence at first enhanced and then overwhelmed the influence of glacioeustasy as part of the region's response to the north European Scandian deformation

Metallurgy of high-silicon steel parts produced using selective laser melting

The metallurgy of high-silicon steel (6.9%wt.Si) processed using Selective Laser Melting (SLM) is presented for the first time in this study. High-silicon steel has great potential as a soft magnetic alloy, but its employment has been limited due to its poor workability. The effect of SLM-processing on the metallurgy of the alloy is investigated in this work using microscopy, X-Ray Diffraction (XRD) and Electron Backscatter Diffraction (EBSD). XRD analysis suggests that the SLM high-silicon steel is a single ferritic phase (solid solution), with no sign of phase ordering. This is expected to have beneficial effects on the material properties, since ordering has been shown to make silicon steels more brittle and electrically conductive. For near-fully dense samples, columnar grains with a high aspect ratio and oriented along the build direction are found. Most importantly, a fibre-texture along the build direction can be changed into a cube-texture when the qualitative shape of the melt-pool is altered (from shallow to deep) by increasing the energy input of the scanning laser. This feature could potentially open the path to the manufacture of three-dimensional grain-oriented high-silicon steels for electromechanical applications

Chitinozoan biostratigraphy of the regional Arenig Series in Wales and correlation with the global Lower–Middle Ordovician series and stages

The Arenig Fawr area of North Wales constitutes the type area for the British Lower to Middle Ordovician Arenig Series and is complemented by sections in the Carmarthen and Whitland areas of South Wales. We describe chitinozoan assemblages from both areas in order to aid correlation of the Arenig Series in its type region with the global Ordovician series and stages. Chitinozoans recorded from Arenig Fawr provide permissive rather than conclusive evidence but suggest that the Henllan Ash Member correlates with the upper Floian Stage Slice Fl3 or lower Dapingian Stage Slice Dp1. Better results were obtained from South Wales where six chitinozoan assemblages are distinguished, ranging in age from late Tremadocian to middle Darriwilian (early Llanvirn). Most species are known from South China, Gondwana and/or Baltica where there are controls on ranges. They show that much of the lower Arenig (Moridunian) succession in South Wales correlates with the upper Floian Stage (Fl3). Correlatives of the lower and middle Floian Stage (Fl1, Fl2), if present, must be represented by the Ogof Hên Formation and lowest Carmarthen Formation. Chitinozoan assemblages from the upper Arenig Series (Fennian Stage) are more readily correlated with Gondwanan biozones and indicate correlation of the Fennian Stage with the Dapingian and lower Darriwilian (Dw1) stages. The middle Arenig Whitlandian Stage is constrained in South Wales to an interval from the uppermost Floian Stage to the basal Dapingian Stage, resulting in an inferred increased rate of sediment accumulation