Crystalline structure,orientation and nucleation of the nacreous tablets in the cephalopod <Emphasis Type="Italic">Nautilus</Emphasis>

The nacreous tablets in the Nautilus shell have similar crystalline structure as the tablets in the gastropod Gibbula shell. Etching with Mutvei’s solution reveals that each tablet is composed of vertical crystalline columns that are structurally similar to the acicular crystallites in the outer spherulitic-prismatic layer of the shell wall. The columns are attached to each other to form numerous vertical crystalline lamellae, oriented parallel to the longitudinal axis of the tablet. It is still unknown whether or not the orientation of the vertical lamellae corresponds to that of the crystallographic a- or b-axis. The orientation of the crystalline lamellae in the adjacent tablets is parallel in some nacreous laminae, but random in other laminae. Similar large variation was found in the nacreous tablets of the gastropod and bivalve shells. The nucleation sites of the nacreous tablets are predominantly situated on the peripheral portion of the upper surface of the preceding tablet, both in the shell wall and septa. The “aragonite-nucleating proteins” in the central portion of the crystal imprints of the organic interlamellar sheets, described by several writers, have therefore a negative correlation with the nucleation sites of the nacreous tablets.     

Skeletal chemistry of Nautilus and its taxonomic significance

The strontium (Sr) and magnesium (Mg) chemistry of the shell wall and septum as well as the spherulitic-prismatic and nacreous layers of these structures was determined for Nautilus species: N. belauensis, N. macromphalus, N. pompilius and N. scrobiculatus. Each species of Nautilus exhibits greater variability and higher concentrations of Mg in juvenile portions of the shell than in more mature portions of the shell. This decrease in the variability and amount of Mg in the aragonite lattice suggests a physiochemical system which becomes more efficient with time relative to carbonate production. Statistically significant differences in the Sr and Mg content of spherulitic-prismatic and nacreous layers of the shell and septum indicate that these layers were formed from extracellular fluids of different compositions. Concentrations of Sr and Mg in aragonite of the shell wall are characteristic for each species and sufficiently invariant within species to allow species of Nautilus to be distinguished statistically on the basis of either the Sr or Mg content of the shell wall or the Mg content of septa.     

Origin of intracameral sheets in ammonoids

The distribution, morphology and mutual relationships of cameral sheets in ammonoids are revised and re-evaluated. Taking into account recent models of ammonoid septum and chamber formation, three different origins can be attributed to the morphological types of sheets: (1) membranes replicated by the rear mantle (pseudosepta and septal linings), (2) membranes secreted sequentially and/or stretched across the chamber (horizontal membranes and chamber linings) and (3) products of desiccation of the cameral liquid (transverse and siphuncular sheets), presumably a cameral hydrogel. Sheets are always preserved near the siphuncular area, because as the cameral liquid was pumped out from the chamber it became progressively richer in dissolved mucus. In the last-formed drops, or menisci, this mucus adhered to the surface of the previously secreted sheets and, on dehydration, it also replicated the surface of the residual reservoirs, producing desiccation sheets. On the basis of the new evidence, changes in the shape of the rear mantle in Triassic ammonoids can be reconstructed. In general, deformations affected the rounded or bottle-neck saddles, which deflated after detachment of the last-formed septum and reinflated when the position of the next septum was reached. The rest of the elements of the septal epithelium were affected to a much lesser extent. One of the functions of those cameral sheets secreted by the rear body was related to a more efficient transport of the cameral liquid upon decoupling from the siphuncular tube. Ammonoids, Triassic, septum, chamber growth, cameral sheets.     

Preferential predatory peeling: Ammonoid vs. nautiloid shells from the Upper Carboniferous of Texas,USA

Narrow groove-like excavations on ammonoid and coiled nautiloid shells are rare in Upper Carboniferous units from Texas, USA. The morphological characteristics of the excavation grooves typically are confined to the ventral and ventrolateral parts of the outer whorl of the shell, are narrower than the length, and have irregular edges where small segments or chips of shells have been removed. Analysis of these features reveals a statistically significant preferential occurrence on ammonoids (1.195% of ca. 3515 specimens) as compared to coiled nautiloids (0.506% of ca. 2965 specimens). The ammonoids typically have longer excavations that penetrate the phragmocone more frequently than those observed in the coiled nautiloids. The groove-like excavations were probably formed by the removal and peeling of shell material by one or more predatory or scavenging arthropods to obtain organic material (tissue and membranes) within the ammonoid and nautiloid body chambers and phragmocones. The excavations probably occurred when the cephalopod was alive (i.e., the cause of death) or shortly after the cephalopod's death. There is no evidence that the excavations are related to sheltering by the excavating organism.     

Molecular cloning and characterization of perlucin from the freshwater pearl mussel, Hyriopsis cumingii

Perlucin is an important functional protein that regulates shell and pearl formation. In this study, we cloned the perlucin gene from the freshwater pearl mussel Hyriopsis cumingii, designated as Hcperlucin. The full-length cDNA transcribed from the Hcperlucin gene was 1460 bp long, encoding a putative signal peptide of 20 amino acids and a mature protein of 141 amino acids. The mature Hcperlucin peptide contained six conserved cysteine residues and a carbohydrate recognition domain, similar to other members of the C-type lectin families. In addition, a “QPS” and an invariant “WND” motif near the C-terminal region were also found, which are extremely important for polysaccharide recognition and calcium binding of lectins. The mRNA of Hcperlucin was constitutively expressed in all tested H. cumingii tissues, with the highest expression levels observed in the mantle, adductor, gill and hemocytes. In situ hybridization was used to detect the presence of Hcperlucin mRNA in the mantle, and the result showed that the mRNA was specifically expressed in the epithelial cells of the dorsal mantle pallial, an area known to express genes involved in the biosynthesis of the nacreous layer of the shell. The significant Hcperlucin mRNA expression was detected on day 14 post shell damage and implantation, suggesting that the Hcperlucin might be an important gene in shell nacreous layer and pearl formation. The change of perlucin expression in pearl sac also confirmed that the mantle transplantation results in a new expression pattern of perlucin genes in pearl sac cells that are required for pearl biomineralization. These findings could help better understanding the function of perlucin in the shell and pearl formation.     

Die Mikro- und Ultrastruktur abgedeckter Hohlelemente und die Conellen des Ammoniten-Gehäuses

Hollow floored spines in the shell ofKosmoceras (Kosmoceras) spinosum (Sow.) and the hollow floored keel ofEleganticeras elegantulum (Young & Bird) have been studied with the scanning electron microscope. In both cases the shell wall is complete in so far as it consists of the outer prismatic layer, the nacreous layer and at least the distal zones of the inner prismatic layer. Both types of hollow shell elements are separated from the lumen of the whorl by a floor which is made up by the proximal zones of the inner prismatic layer. This explains why conellae occur, with preference, along the floors of hollow spines and keels. The origin of primary aragonitic conellae and of secondary calcitic conellae is discussed as well as their dependence on structural properties of the corresponding shell layer, which is the inner prismatic layer. An attempt is made to reconstruct the way of formation of the floored hollow spines and the floored hollow keels by the mantle epithelium.     

Analysis of a Carboniferous embryonic ammonoid assemblage implications for ammonoid embryology

An embryonic ammonoid assemblage was discovered in a carbonate concretion recovered from a dysoxic, relatively offshore marine shale of Virgilian (Upper Pennsylvanian) age in Kansas, USA. The assemblage consists primarily of two species of the Goniatitina, Aristocerassp. and Vidrioceras sp., whose initial chambers (protoconchs) differ in size and shape. Microscopic observations of serial thin sections of specimens at different growth stages reveal the sequence of embryonic shell development starting with the formation of the initial chamber and ending with the synchronous secretion of a prismatic proseptum and nacreous swelliig (primary varix) at the aperture. The mode of occurrence of the embryonic shells of the two species in the concretion suggests that these ammonoids produced numerous small offspring, a reproductive strategy similar to that in many extant coleoids. □ Ammonoids, embryonic shells, development, Carboniferous, Kansas.     

Morphological studies on the calcification process in the fresh-water mussel Amblema

Light microscopy, transmission electron microscopy, scanning electron microscopy, various histochemical procedures for the localization of mineral ions, and analytical electron microscopy have been used to investigate the mechanisms inherent at the mantle edge for shell formation and growth in Amblema plicata perplicata, Conrad. The multilayered periostracum, its component laminae formed from the epithelia lining either the periostracal groove or the outer mantle epithelium (of the periostracal cul de sac), appears to play the major regulatory and organizational role in the formation of the component mineralized layers of the shell. Thus, the inner layer of the periostracum traps and binds calcium and subsequently gives rise to matricial proteinaceous fibrils or lamellar extensions which serve as nucleation templates for the formation and orientation of the crystalline subunits (rhombs) in the forming nacreous layer. Simultaneously, the middle periostracal layer furnishes or provides the total ionic calcium pool and the matricial organization necessary for the production of the spherical subunits which pack the matricial ‘bags’ of the developing prismatic layer. The outer periostracal layer appears to be a supportive structure, possibly responsible for the mechanical deformations which occur in the other laminae of the periostracum. The functional differences in the various layers of the periostracum are related to peculiar morphological variables (foliations, vacuolizations, columns) inherent in the structure and course of this heterogeneous (morphologically and biochemically) unit. From this study, using the dynamic mantle edge as a morphological model system, we have been able to identify at least six interrelated events which culminate in the production of the mature mineralized shell layers (nacre, prisms) at the growing edge of this fresh-water mussel.     

A new model for periostracum and shell formation in Unionidae (Bivalvia, Mollusca)

The periostracum in Unionidae consists of two layers. The outer one is secreted within the periostracal groove, while the inner layer is secreted by the epithelium of the outer mantle fold. The periostracum reaches its maximum thickness at the shell edge, where it reflects onto the shell surface. Biomineralization begins within the inner periostracum as fibrous spheruliths, which grow towards the shell interior, coalesce and compete mutually, originating the aragonitic outer prismatic shell layer. Prisms are fibrous polycrystalline aggregates. Internal growth lines indicate that their growth front is limited by the mantle surface. Transition to nacre is gradual. The first nacreous tablets grow by epitaxy onto the distal ends of prism fibres. Later growth proceeds onto previously deposited tablets. Our model involves two alternative stages. During active shell secretion, the mantle edge extends to fill the extrapallial space and the periostracal conveyor belt switches on, with the consequential secretion of periostracum and shell. During periods of inactivity, only the outer periostracum is secreted; this forms folds at the exit of the periostracal groove, leaving high-rank growth lines. Layers of inner periostracum are added occasionally to the shell interior during prolonged periods of inactivity in which the mantle is retracted.     

PERNICIÖSE EPÖKIE VON PLACUNOPSIS AUF CERATITES

An adult Ceratites semipartitus encrusted by epifaunal Placunopsis ostracina is described. The epizoans are orientated in relation to the slope of the substratum and according to their time of attachment to the ammonoid shell. Geometrical analysis of the orientation leads to the following conclusions: (1) During and after the growth of the last whorl, several, probably 4, swarms of larvae of Placunopsis settled upon the Ceratites specimen. (2) The dead body was for some time kept upright, the living chamber showing upwards. During this time, larvae of Placunopsis became attached again. (3) The ammonoid shell was then laid down, its left flank in contact with the sediment. In this position it was for the last time settled upon by Placunopsis which died whilst juvenile.
Provided that the phases of attachment corresponded to annual spawning periods of Placunopsis , the last whorl of the Ceratites was formed with decreasing velocity of growth within about 4 years.
Placunopsis became attached with its right valve, the posterior end of the body showing upwards, and the direction of growth being declined from the vertical by 65°. Placunopsis either preferred to settle within a distinct zone of the living chamber of the ammonoid close to the lowermost part of the shell or was able to grow up here only. Ceratites semipartitus was therefore probably nectonic.
Both the host and the epizoan were influenced to their disadvantage by their fatal partnership. A relationship of this kind therefore should not be called a symbiosis.     

Studies on chitin and calcification in the inner layers of the shell of Anodonta cygnea

Summary An orthorhombic structure -chitin, probably in the form of a chitin-protein complex, was identified in the matrix of the shell of Anodonta cygnea by X-ray diffraction. Aragonite crystals of pseudohexagonal symmetry were also found by a Lauegram on the nacreous layer of the shell. The orthorhombic structure of these two compounds together with the identical reticular spacing d₁₁₀ corroborate, in Anodonta cygnea, the indirect chitin-aragonite relationships already suggested for molluscan shells.Observations with SEM in the inner surface of the shell showed CaCO₃ crystals with irregular geometrical shapes in spring and summer and regular geometrical shapes in autumn and winter. The more elaborate aspect appearing in winter corresponds to an accurate hexagonal shape. This suggests that the observed variability may depend on the balance between calcium and hydrogen ions in the extrapallial fluid.Abbreviations OME outer mantle epithelium - SEM scanning electron microscopy     

Desmoceras (Pseudouhligella) intrapunctatum n. sp. (Ammonoidea) aus dem Unter-Albium von NW-Madagaskar mit Ritzstreifen

The new ammonoid speciesDesmoceras (Pseudouhligella) intrapunctatum (DesmoceratoideaZittel, 1895) is described and figured from the dark glauconitic marls of the Lower Albian of Ambatolafia (Mahajanga Basin of northwestern Madagascar). On the molds of its body chamber a well developed system of dotted lines is visible originally described from Palaeozoic ammonoids and nautiloids (= ?Ritzstreifen“Sandberger & Sandberger 1850 sensuTozer 1972). These internal shell structures have been originated by small ridges of the inner prismatic layer as the result of an incomplete mineralization. “Ritzstreifen” could not have observed in any other ammonoid species accompanied withDesmoceras (Pseudouhligella) intrapunctatum, despite of the same excellent aragonitic shell preservation. Therefore, this unique character is interpreted as taxonomically significant on species level.     

The Evolution and Development of Cephalopod Chambers and Their Shape

The Ammonoidea is a group of extinct cephalopods ideal to study evolution through deep time. The evolution of the planispiral shell and complexly folded septa in ammonoids has been thought to have increased the functional surface area of the chambers permitting enhanced metabolic functions such as: chamber emptying, rate of mineralization and increased growth rates throughout ontogeny. Using nano-computed tomography and synchrotron radiation based micro-computed tomography, we present the first study of ontogenetic changes in surface area to volume ratios in the phragmocone chambers of several phylogenetically distant ammonoids and extant cephalopods. Contrary to the initial hypothesis, ammonoids do not possess a persistently high relative chamber surface area. Instead, the functional surface area of the chambers is higher in earliest ontogeny when compared to Spirula spirula. The higher the functional surface area the quicker the potential emptying rate of the chamber; quicker chamber emptying rates would theoretically permit faster growth. This is supported by the persistently higher siphuncular surface area to chamber volume ratio we collected for the ammonite Amauroceras sp. compared to either S. spirula or nautilids. We demonstrate that the curvature of the surface of the chamber increases with greater septal complexity increasing the potential refilling rates. We further show a unique relationship between ammonoid chamber shape and size that does not exist in S. spirula or nautilids. This view of chamber function also has implications for the evolution of the internal shell of coleoids, relating this event to the decoupling of soft-body growth and shell growth.     

SOME LESSER KNOWN FEATURES OF THE ANCIENT CEPHALOPOD ORDER ELLESMEROCERIDA (NAUTILOIDEA, CEPHALOPODA)

Abstract:  Three specimens of the small breviconic ellesmeroceratid Paradakeoceras minor Flower, 1964 from the Tremadocian of the New York area preserve the annular elevation and muscle scars in moulds of the body chamber. The annular elevation is positioned at the base of the body chamber and is wider on the convex side of the shell than on the concave side. Multiple paired muscle scars can be seen within this annular elevation. A well-preserved body chamber of the breviconic ellesmeroceratid Levisoceras cf. edwardsi Ulrich, Foerste and Miller is described. Its body chamber shows a strong anterior–posterior asymmetry, which is common within the Ellesmeroceratida. The shape of the body chamber and of the soft body attachment structures has led to a reconstruction of an ellesmeroceratid soft body that is organized like a primitive conchiferan mollusc. Based on this reconstruction, a tryblidian cephalopod ancestor is supported. An evolutionary scenario is reconstructed from an ancestral nautiloid that is stretched along the anterior–posterior axis, and has serially arranged shell muscles and a small mantle cavity, towards a modern cephalopod with a dorsal–ventral body orientation, reduced number of shell muscles and a large mantle cavity.     

Structure and composition of the septal nacreous layer of Nautilus macromphalus L. (Mollusca, Cephalopoda)

The nacreous layer of Mollusca is the best-known aragonitic structure and is the usual model for biomineralization. However, data are based on less than 10 species. In situ observations of the septal nacreous layer of the cephalopod Nautilus shell has revealed that the tablets are composed of acicular laths. These laths are composed of round nanograins surrounded by an organic sheet. No hole has been observed in the decalcified interlamellar membranes. A set of combined analytical data shows that the organic matrices extracted from the nacreous layer are glycoproteins. In both soluble and insoluble matrices, S amino acids are rare and the soluble organic matrices have a higher sulfated sugar content than the insoluble matrices. It is possible that the observed differences in the structure and composition of the nacreous layers of the outer wall and septa of the Nautilus shell have a dual origin: evolution and functional adaptation. However, we have no appropriate data as yet to answer this question.     

Palaeobiology and taxonomy ofPachyperna laverdana Oppenheim,an Eocene bivalve of Mesozoic heritage

Pachyperna laverdana is a large Eocene bivalve characterized by an extremely thick shell wall. Rediscovery of the type locality(Pernabank Auctt.) after more than a century has made it possible to collect abundant material which is used here to provide a better morphological definition of the taxon. In particular, indication is given of its broad intraspecific variability mostly due to the gregareous habit (ecomorphism) and by change of mode of life through ontogenesis. As regards the latter factor, functional analysis of the shell suggests that in its early juvenile stages the bivalve was an epibyssate, pleurothetic form, attached to hard substrata. Then, it moved to soft-bottom substrates, where a “heavy-weight” strategy was developed to compensate for a weak byssal attachment. In the adult stage, it may be considered a reclining, orthothetic form. The shell is made up of a thin outer layer formed by simple prismatic calcite and by thick, aragonitic, irregular fibrous-prismatic inner layers, both with a well marked periodicity of growth. Mechanical, functional and systematic significance of shell microstructures are discussed. The diagnosis of the genusPachyperna is herewith emended.     

Aragonitic dendritic prismatic shell microstructure in Thracia (Bivalvia,Anomalodesmata)

The shells of most anomalodesmatan bivalves are composed of an outer aragonitic layer of either granular or columnar prismatic microstructure, and an inner layer of nacre. The Thraciidae is one of the few anomalodesmatan families whose members lack nacreous layers. In particular, shells of members of the genus Thracia are exceptional in their possession of a very distinctive but previously unreported microstructure, which we term herein “dendritic prisms.” Dendritic prisms consist of slender fibers of aragonite which radiate perpendicular to, and which stack along, the axis of the prism. Here we used scanning and transmission electron microscopical investigation of the periostracum, mantle, and shells of three species of Thracia to reconstruct the mode of shell calcification and to unravel the crystallography of the dendritic units. The periostracum is composed of an outer dark layer and an inner translucent layer. During the free periostracum phase the dark layer grows at the expense of the translucent layer, but at the position of the shell edge, the translucent layer mineralizes with the units typical of the dendritic prismatic layer. Within each unit, the c‐axis is oriented along the prismatic axis, whereas the a‐axis of aragonite runs parallel to the long axis of the fibers. The six‐rayed alignment of the latter implies that prisms are formed by {110} polycyclically twinned crystals. We conclude that, despite its distinctive appearance, the dendritic prismatic layer of the shell of Thracia spp. is homologous to the outer granular prismatic or prismatic layer of other anomalodesmatans, while the nacreous layer present in most anomalodesmatans has been suppressed.