Ostracodes swallowed by Palaeozoic corals?

Middle Devonian heliolitids and favositids from Central Bohemia, belonging to Heliolites 'intermedius' LeMaitre and Favosites goldfussi Orbigny , incorporated ostracode shells within their living corallite structures. The ostracode shells were sealed in by skeletal tissue that was septal in origin (Heliolites) or they were roofed over by tabulae (Favosites). The foreign shell was near the axis of the polyp when trapped within the coral skeleton. Only ostracodes, not other rounded shells or sedimentary particles, were trapped in this way. Approximately one in 30 favositid corallites and one in 70 heliolitid corallites display this peculiar condition, where the ostracode shells seem to have been swallowed by the polyps. A probable scenario involves the injury of the mouth area and the trapping of the ostracodes. A high probability that the basal part of the polyp experienced a controlled penetration is the most striking part of the process. □ Favositids, heliolitids, ostracodes, coral growth violence, behavior, Middle Devonian, Bohemia.     

Ultrastructural features and elemental distribution in eggshell during pre and post hatching periods in the green turtle, Chelonia mydas at Ras Al-Hadd, Oman

Eggshells were randomly collected from turtle nests immediately after oviposition and at the end of incubation to examine the ultrastructural features using scanning JSM-5600LV microscopy. Three layers were recognized; an outer calcareous, a middle multistrata and an inner membrane. The calcareous layer had loose nodular units varying in shape and size without interlocking attachments. In freshly laid eggs, each nodular unit had spicules arranged in folded stacks. The spicules became unfolded during incubation, to form radiating configurations. Elemental composition and mapping of the layers were analyzed using energy dispersive spectroscopy (EDS). The elements were unevenly distributed throughout the eggshell and Ca²⁺ decreased significantly after hatching. X-ray diffraction was used to identify the crystals of the eggshells. It revealed that nodular units of the calcareous were made up of CaCO₃, as aragonite (91%), calcite (6%) and vaterite (3%). The middle layer consisted of organic amorphous material with aragonite (89%) and calcite (11%). The shell membrane consisted of reticular fibers with crystals predominantly of NaCl halite. Thermogravimetry analysis of the calcareous layer indicated a complete evaporation of bonded H₂O at 480 °C and CO₂ at 830 °C. Using the differential thermal analysis (DTA), aragonite was transformed to stable calcite at 425 °C.     

CRYSTAL PROPERTY OF THE LARVAL SEA URCHIN SPICULE*

A pair of pluteus skeletal spicules arises from a pair of calcareous granules via the triradiate form. In polarized light, each spicule behaves as though carved out of a single crystal of magnesian calcite. The optic axis lies perpendicular to the plane of the triradiate and parallel to the body rod of the pluteus. However, in the scanning electron microscope, the spicule surface appeared smooth or somewhat spongy and manifested no crystal faces. Neither etching nor fracturing revealed underlying crystalline texture. Nevertheless, rhombohedral calcite crystals could be grown epitaxially onto isolated spicules immersed in a medium containing CaCl₂ and NaHCO₃. The optic axes of all crystals coincided with the optic axis of the spicule on which they were grown. Corresponding faces of the crystals were all aligned parallel to each other despite the complex shape of each spicule. Where the left and right spicules joined, two mutually tilted sets of crystals were observed but not crystals of intermediate orientation. Thus, the sea urchin larval spicule is built from a stack of molecularly contiguous microcrystals but its overall shape is generated by the mesenchyme cells independent of the magnesian calcite crystal habit.     

Where's the glass? Biomarkers,molecular clocks,and microRNAs suggest a 200‐Myr missing Precambrian fossil record of siliceous sponge spicules

The earliest evidence for animal life comes from the fossil record of 24-isopropylcholestane, a sterane found in Cryogenian deposits, and whose precursors are found in modern demosponges, but not choanoflagellates, calcareans, hexactinellids, or eumetazoans. However, many modern demosponges are also characterized by the presence of siliceous spicules, and there are no convincing demosponge spicules in strata older than the Cambrian. This temporal disparity highlights a problem with our understanding of the Precambrian fossil record – either these supposed demosponge-specific biomarkers were derived from the sterols of some other organism and are simply retained in modern demosponges, or spicules do not primitively characterize crown-group demosponges. Resolving this issue requires resolving the phylogenetic placement of another group of sponges, the hexactinellids, which not only make a spicule thought to be homologous to the spicules of demosponges, but also make their first appearance near the Precambrian/Cambrian boundary. Using two independent analytical approaches and data sets – traditional molecular phylogenetic analyses and the presence or absence of specific microRNA genes – we show that demosponges are monophyletic, and that hexactinellids are their sister group (together forming the Silicea). Thus, spicules must have evolved before the last common ancestor of all living siliceans, suggesting the presence of a significant gap in the silicean spicule fossil record. Molecular divergence estimates date the origin of this last common ancestor well within the Cryogenian, consistent with the biomarker record, and strongly suggests that siliceous spicules were present during the Precambrian but were not preserved.     

Biomineralization of the spicules of sea urchin embryos

The formation of calcareous skeletal elements by various echinoderms, especially sea urchins, offers a splendid opportunity to learn more about some processes involved in the formation of biominerals. The spicules of larvae of euechinoids have been the focus of considerable work, including their developmental origins. The spicules are composed of a single optical crystal of high magnesium calcite and variable amounts of amorphous calcium carbonate. Occluded within the spicule is a proteinaceous matrix, most of which is soluble; this matrix constitutes about 0.1% of the mass. The spicules are also enclosed by an extracellular matrix and are almost completely surrounded by cytoplasmic cords. The spicules are deposited by primary mesenchyme cells (PMCs), which accumulate calcium and secrete calcium carbonate. A number of proteins specific, or highly enriched, in PMCs, have been cloned and studied. Recent work supports the hypothesis that proteins found in the extracellular matrix of the spicule are important for biomineralization.     

Halysitid tabulates: sponges in corals' clothing

Abundant pyritic pseudomorphs of monaxonic siliceous spicules (ophirhabds and ?heloclones) have been found entrapped in the calcareous skeleton of the halysitid tabulate Quepora ?agglomeratiformis (Whitfield) from late Ordovician limestones of Frobisher Bay, Baffin Island, Canada. The finding indicates a poriferan (choristid or sublithistid) affinity of halysitids, early Palaeozoic marine fossils related so far to corals. They probably derived from a monaxonic group of early demosponges that adapted during the Ordovician to Ca²⁺ stress conditions in epicontinental seas by excreting the excessive Ca²⁺ influx to their tissues as variously designed chains of basally secreted calcareous tubes.     

苏北盆地阜宁组钙质超微化石由来初探

对苏北盆地4个钻孔300余块心样品进行了分析,在阜宁组的8块样品中发现了钙质超微化石,计3属1种及3个比较种,化石属种单调,保存较 ,受到严重的溶蚀或增生,结合这些种类的分布时代和产出层位,很难排除苏北盆地阜宁组钙质超微化石再沉积的属性。     

Spicule form in calcareous sponges (Porifera: Calcarea). The principle of uniplanar curvature

A study of the shapes of secondary sagittal tri- and quadri-radiate spicules of the calcareous sponge Leuconia fistulosa has revealed that the paired rays appear kinked, with straight sections in between the kinks, when the spicules are viewed along the optic axis of the constituent calcite. The secondary sagittal spicules thus conform with monaxons and with per-regular and primary sagittal tri- and quadri-radiates, in general, in that whenever a ray is curved, the curvature is always in a single plane that includes the optic axis. The kinks are not necessarily distributed in symmetrical fashion along the paired rays. A mechanism is proposed whereby uniplanar curvature could be achieved.     

Nano-cluster composite structure of calcitic sponge spicules--a case study of basic characteristics of biominerals

Spicules of calcareous sponges are elaborately shaped skeletal elements that nonetheless show characteristics of calcite single-crystals. Our atomic force microscopic and transmission electron microscopic investigation of the triradiate spicules of the sponge Pericharax heteroraphis reveals a nano-cluster structure with mostly well-aligned small crystal domains and pockets with accumulated domain misalignments. Combined high-resolution and energy-filtering transmission electron microscopy revealed carbon enrichments located in between crystal domain boundaries, which strongly suggests an intercalated network-like proteinaceous organic matrix. This matrix is proposed to be involved in the nano-clustered calcite precipitation via a transient phase that may enable a 'brick-by-brick' formation of composite and yet single-crystalline spicules with elaborate morphologies. This composite cluster structure reduces the brittleness of the material by dissipating strain energy and deflecting crack propagation from the calcite cleavage planes, but the lattice symmetry and anisotropic growth properties of calcite still play a major role in the morphogenesis of these unusual calcite single-crystals. Our structural, crystallographic, textural, and chemical analysis of sponge spicules corroborates the view that nano-clustered crystal growth, induced by organic matrices, is a basic characteristic of biomineralisation that enables the production of composite materials with elaborate morphologies.     

Matrix and Mineral in the Sea Urchin Larval Skeleton

The endoskeletal spicules of sea urchin larvae are composed of calcite, a surrounding extracellular matrix, and small amounts of occluded matrix proteins. The spicules are formed by primary mesenchyme cells (PMCs) in the blastocoel of the embryo, where they adopt stereotypical locations, thereby specifying where spicules will form. PMCs also fuse to form cytoplasmic cords connecting the cell bodies, and it is within the cords that spicules arise. The mineral phase contains 5% Mg as well as Ca, and about 0.1% of the mass is protein. The matrix and mineral form concentric plies, and the composite has different physical properties than those of pure calcite. The calcite diffracts as a single crystal and is composed of well-ordered, but not perfectly ordered, microdomains. There is evidence for adsorption of matrix proteins to specific crystal faces at domain boundaries, which may help regulate crystal growth and texture. Immature spicules contain considerable precipitated amorphous CaCO3, and PMCs also have vesicles that contain amorphous CaCO3. This suggests the hypothesis that the cellular precursor to the spicules is actually amorphous CaCO3 stabilized in the cell by protein. The spicule s enveloped by the PMC cord, but is topologically exterior to the cell. The PMC plasmalemma is tightly applied to the developing spicules, except perhaps at the elongating tip. The characteristics, localization, and possible function of the four identified matrix proteins are discussed. SM50, SM37, and PM27 all primarily enclose the mineral, though small amounts are occluded. SM30 is found in cellular vesicles and is probably the principal occluded protein of the spicule.     

KATBERGIA GEN. NOV., A NEW TRACE FOSSIL FROM UPPER PERMIAN AND LOWER TRIASSIC ROCKS OF THE KAROO BASIN: IMPLICATIONS FOR PALAEOENVIRONMENTAL CONDITIONS AT THE P/TR EXTINCTION EVENT

Abstract:  A new ichnogenus and ichnospecies of burrow, Katbergia carltonichnus , are described from Upper Permian and Lower Triassic rocks of the Karoo Basin, South Africa, where they are preserved in pedogenically modified overbank deposits that are interpreted as inceptisols subsequently gleysol overprinted. Sigmoidal burrows consist of a long (≥0.5 m) cylindrical tube, ranging from 1–2 cm in diameter, terminating in a slightly larger living chamber. The burrows are unlined and passively filled, preserving a hierarchy of scratch patterns on the burrow walls. Scratch patterns include longitudinal, transverse, and crescent-shaped markings found around the circumference of the burrow, but which are less densely concentrated on the burrow floor. Calcareous concretions are associated with burrowed siltstone intervals, generally restricted to the lowermost decimetre, with nodules nucleating around burrows. Stable δ¹³C and δ¹⁸O isotope data on calcite cement in the burrow fill, entombing siltstone, and associated calcareous nodules all cluster together when plotted, indicating that nodule formation occurred following burrow horizon abandonment and a rise in regional water table. Isotopic data reflect calcite precipitation under a semi-closed system in saturated conditions. A model for burrow emplacement, abandonment and infill, and subsequent cementation by calcite is presented demonstrating that previous interpretations of Late Permian and Early Triassic palaeosol types associated with the P/Tr extinction event must be re-evaluated.     

Palaeozoic calcareous plankton: evidence from the Silurian of Gotland

In order to investigate whether or not equivalents to modern calcareous plankton existed in Palaeozoic times, extremely well-preserved successions have to be investigated. The Silurian strata on Gotland (Sweden) are exceptionally well preserved because they have not experienced deep burial conditions and tectonic stress, due to their position on the stable Baltic Shield. Scanning electron microscope investigations of polished, slightly etched rock surfaces revealed the presence of a variety of calcareous micro- and nannofossils. Among these organisms, many can be termed 'calcispheres' (60–100 µm in diameter), whereas others due to their size range (nannofossils) are informally termed herein as 'nannospheres' (10–25 µm in diameter). The systematic attribution of these fossils is unknown ( incertae sedis ). Mesozoic calcispheres are usually attributed to calcareous cysts of dinoflagellates following comparisons of ultrastructure with modern species. The abundance of different calcispheres in the Silurian sediments of Gotland and the observation that many of the calcareous microfossils occur in distinctly different facies as well as their spherical shape indicate that they probably belong to calcareous micro- and nannoplankton. We therefore conclude that calcareous plankton most probably existed already during the Palaeozoic, but can only be observed under conditions of exceptional preservation.     

Triassic/Jurassic carbonates from the Hochfelln Mountain (Northern Calcareous Alps)—its facies, silicified fauna and implications for the end-Triassic biotic crisis

A suite of Early Mesozoic (Late Triassic, Norian to Early Jurassic) calcareous beds was studied from the Hochfelln Mountain in the Northern Calcareous Alps (NCA, South Germany). The Hauptdolomit Group consists of thick peritidal deposits and is overlain by basin deposits of the Rhaetian Kössen Formation and Rhaetian reefoidal limestone with corals. Unlike many other sections in the Tethys realm, coral growth seems to continue into the Jurassic or starts again relatively early within the Early Jurassic. Silicified corals and other marine invertebrates are present in the calcareous, micritic Hochfelln Beds. A re-examination of previously collected ammonite material indicates the presence of Coroniceras sp. which suggests an Early Sinemurian age for the Hochfelln Beds. Abundant sponge spicules (spiculites) suggest that sponges were the source for the silicification. The site produced one of the most diverse Early Jurassic (Sinemurian) gastropod faunas of the NCA (25–30 species, some undescribed). The relatively diverse Early Sinemurian gastropod fauna and coral growth indicate rapid recovery from the end-Triassic biotic crisis.     

Phylogenetic comparison of spicule networks in cryptobranchiate dorid nudibranchs (Gastropoda, Euthyneura, Nudibranchia, Doridina)

Many dorid nudibranchs possess large numbers of calcareous spicules in their mantle, gill, rhinophores and foot. However, the arrangements of these structures and their differences among taxa are poorly known. Spicule networks were stained with Alizarin red and compared among 12 species of cryptobranchiate dorid nudibranchs and four outgroups. Three general types of networks were found: a cobweb-like, unbraced framework of one or few spicules per side; a ramifying system of thick, spiculated tracts; and a lattice-like arrangement of distinct radial and circumferential tracts. The Discodorididae species investigated shared a cobweb-like network and papillae supported by a ring of spicules, while the Porostomata showed consistent characters leading to a lattice-like network with larger spicules in the central notum. The Dorididae studied were not cohesive, but each species shared characters with the aforementioned groups. Therefore, spicule network form may provide new characters to help resolve the phylogeny of Doridina.     

The localization of occluded matrix proteins in calcareous spicules of sea urchin larvae

The sea urchin embryo forms calcareous endoskeletal spicules composed of calcite and an occluded protein matrix. Though the latter is approximately 0.1% of of the mass, the composite has substantially altered material properties, e.g., conchoidal fracture planes and increased hardness. Experiments were conducted to examine the localization of matrix proteins occluded in the mineral by use of immunocytochemistry coupled with scanning electron microscopy (SEM). The isolated, unfixed spicules were etched under relatively gentle conditions and exposed to affinity purified antibodies made against two different matrix proteins, as well as an antibody to the entire constellation of matrix proteins. Immunogold tagged secondary antibody was used to observe antibody localization in the back scatter mode of SEM. All proteins examined were very widely distributed throughout the calcite, supporting a model of the structure in which a multiprotein assemblage is woven with fine texture around microcrystalline domains of calcite. Gentle etching revealed a laminar arrangement of calcite solubility, consistent with a stepwise deposition of matrix and mineral to increase girth of the spicule.     

Epizoic bryozoans and corals as indicators of life and post-mortem orientations of the Devonian brachiopod Meristella

The in-life and post-mortem orientations of the Lower Devonian brachiopod Meristella atoka from the Haragan Formation (Lower Devonian; south-central Oklahoma) are inferred from the distribution of epizoic bryozoans and the orientations of base plates of epizoic corals. Three bryozoans, Cyphotrypa corrugata, Fistuliporella maynardi and Leioclema pulchellum, and one coral, Favosites conicus, are considered. Most zoaria that contact the commissure terminate at the commissurc, and a few zoaria terminate at growth lines. This suggests that the bryozoans were primarily life associates of Meristella atoka. Collectively, these three bryozoan species most extensively encrusted marginal sectors of the brachial valve and are very rare on the posteromedian sector (the umbo) of the pedicle valve. This distributional pattern indicates that the preferred living orientation of Meristella atoka was umbo-down (posteromedian sector of the pedicle valve resting on or buried in the substratum) with the commissure steeply inclined to the sediment water interface. Most coralla of Favosites conicus that contacted the commissure encrusted over the commissure. This indicates that Favosites conicus either preferentially encrusted Meristella atoka post-mortem or colonized living brachiopods but subsequently caused them to die. Furthermore, Favosites conicus most extensively encrusted anterior sectors of the brachial valve, especially the fold. The lateral and anterior orientations of the commissure with respect to the base plates (holothecae) of Favosites conicus indicates that the brachiopods were oriented approximately horizontal with respect to the base plates. This suggests that the preferred post-mortem orientation of Meristella atoka was resting nearly horizontally on the substratum. These data and interpretations confirm previously inferred in-life orientations of Meristella atoka and are consistent with post-mortem orientations hypothesized for elongate-oval athyrid brachiopods. □Brachiopoda, Athyridacea, Meristella atoka , palaeoecology, autecology, epizoans, bryozoans, corals, sedimentology, Lower Devonian, North America, Oklahoma.     

Danian cold‐water corals from the Baunekule facies,Faxe Formation,Denmark: a rare taphonomic window of a coral mound flank habitat

Well‐preserved cold‐water corals are comparatively rare in the fossil record. This is partly due to the very low fossilization potential of the predominantly aragonitic corals but also due to the fact that coral ecosystems of deep water are a geologically young development. A Middle Danian cold‐water coral mound complex is well exposed in Faxe Quarry, Denmark. The coral mounds are intercalated with bryozoan mounds of various sizes and form the Faxe Formation. The coral limestone displays large variations in diagenesis, and this complicates the palaeoecological reconstructions. However, the Baunekule facies from the Faxe Formation contain a well‐preserved originally aragonitic and calcitic fauna. The aragonitic skeletons have been recrystallized to calcite during early diagenesis and the excellent preservation makes taxonomic identifications straightforward. A diverse fauna of ten scleractinian coral species, nine stylasterine coral species and seven octocoral species has been described from the Baunekule facies. The fossil fauna represents an ecological niche between the dead coral framework and coral rubble on a flank of a growing Dendrophyllia coral mound with multiple colonization events. The diversity and relative abundance of the fossil scleractinian corals are comparable to the modern settings in the NE Atlantic and Mediterranean. The distribution and diversity of the octocorals and the stylasterine corals are suggested to represent coral gardens as described from modern setting in the NE Pacific. The presence of a diverse and abundant stylasterine fauna suggests a stable palaeoenvironment, probably in a bathymetric depth range of 200–400 metre.     

从底栖生物化石再释贵州独山泥盆系龙洞水段的时代

贵州独山的猴儿山组龙洞水段是一套浅海相碳酸盐沉积,盛产底栖牛物化石.通过对该段地层中所含四射珊瑚、床板珊瑚和腕足类等化石属种的详尽分析,再次证实龙洞水段的地质时代仍应属中泥盆世早期(Eifelian).而不是像某些根据痕迹化石或层序地层学的研究那样将它置于下泥盆统.     

Evidence for spicule homology in calcareous and siliceous sponges: biminerallic spicules in Lenica sp. from the Early Cambrian of South China

Botting, J.P., Muir, L.A., Xiao, S., Li, X. & Lin, J.‐P. 2012: Evidence for spicule homology in calcareous and siliceous sponges: biminerallic spicules in Lenica sp. from the Early Cambrian of South China. Lethaia, Vol. 45, pp. 463–475. The relationships of the extant sponge classes, and the nature of the last common ancestor of all sponges, are currently unclear. Early sponges preserved in the fossil record differ greatly from extant taxa, and therefore information from the fossil record is critical for testing hypotheses of sponge phylogenetic relationships that are based on modern taxa. New specimens of the enigmatic sponge Lenica sp., from the Early Cambrian Hetang Biota of South China, exhibit an unusual spicule structure. Each spicule consists of a siliceous core with an axial canal, an organic outer layer and a middle layer interpreted to have been originally calcium carbonate. This finding confirms previous work suggesting the existence of biminerallic spicules in early sponges. Combined with data from other early sponges, the new findings imply that the two fundamental spicule structures of modern sponges were derived from a compound, biminerallic precursor. Spicules are therefore homologous structures in Calcarea and Silicea, and if sponges are paraphyletic with respect to Eumetazoa, then spicules may also have been a primitive feature of Metazoa. □Calcarea, Early Cambrian, Hetang Biota, phylogeny, Silicea, taphonomy.