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.     

CONVERGENT SHELL MORPHOLOGY IN INTERTIDAL GASTROPODS

The functional morphology of shell infrastructure in 2 speciesof intertidal trochid was compared with that in 2 species ofnerite. The shell of Monodonta constrictais typical of the majorityof trochids. The shell is composed of 4 layers: a distal layer(calcite), anouter prismatic layer (aragonite), a nacreous layer(aragonite), and an oblique prismatic layer (aragonite). Monodontalabio lacks a distal layer and an oblique prismatic layer. Theoblique prismatic layer is replaced by an inner prismatic layerwhich forms an apertural ridge as a result of deposition andresorption. The shells of Nerita versicolor and N. tessellataconsistof 3 layers: an outer prismatic layer (calcite), a crossedlamellar layer (aragonite), and a complex crossed lamellar layer(aragonite). The complex crossed lamellar layer is covered withinclined platelets which superficially resemble the surfaceof the ique prismatic layer of trochids. In addition, the complexcrossed lamellar layer forms an apertural ridge which is similarin appearance to that of Monodonta labio. The outer surfaceof the mantle of Nerita versicolor and N. tessellata is throwninto a series of large folds which lie in contact with the inclinedplatelets of the omplex crossed lamellar layer. The interactionof the mantle folds with the inclined platelets is thought toserve as a rachet mechanism to aid in extension of themantle;a similar function has previously been proposed for trochids.The apertural ridges of Monodonta labio and Nerita are thoughtto prevent excessive desiccation when these gastropodsare exposedat low tide. ¹Contribution No. 56 of the Tallahassee, Sopchoppy & GulfCoast Marine Biological Association (Received 6 July 1979;     

Microstructure and crystallographic texture of Charonia lampas lampas shell

Charonia lampas lampas shell is studied using scanning electron microscopy and X-ray diffraction combined analysis of the preferred orientations and cell parameters. The Charonia shell is composed of three crossed lamellar layers of biogenic aragonite. The outer layer exhibits a 0 0 1 fibre texture, the intermediate crossed lamellar layer is radial with a split of its c-axis and single twin pattern of its a-axis, and the inner layer is comarginal with split c-axis and double twinning. A lost of texture strength is quantified from the inner layer outward. Unit-cell refinements evidence the intercrystalline organic influence on the aragonite unit-cell parameters anisotropic distortion and volume changes in the three layers. The simulation of the macroscopic elastic tensors of the mineral part of the three layers, from texture data, reveals an optimisation of the elastic coefficient to compression and shear in all directions of the shell as an overall.     

Shell microstructure in the Permian nonmarine bivalve Palaeomutela Amalitzky: Revision of the generic diagnosis

The microstructure of aragonitic and calcitic shells of the genus Palaeomutela Amalitzky, 1891 is examined. The aragonitic shell consists of three main layers, each is distinguished by certain crossed lamellar microstructure: comarginal, radial, and complex. As aragonite is recrystallized into pelitic calcite, microstructural shell features are preserved. Many species of Palaeomutela from localities of different age display the same microstructural pattern, which is possible to regard as a character of generic rank.     

Endocochliate embryo model in the Mesozoic Ammonitida

An endocochliate embryo model for the Mesozoic Ammonitida is proposed based on scanning electron microscopy of the ammonitella (= embryonic shell) stage of well‐preserved Japanese Cretaceous specimens belonging to nine species of five superfamilies. As in other specimens described previously, the ammonitella wall succeeding from the initial chamber ("protoconch") in the species examined consists of the inner prismatic, middle subprismatic and outer prismatic layers, with minute tubercles resting on the outer. Developmental patterns of these structures and their comparison with primary shell formation in modern Nautilus and Spirula suggest that the outer thin prismatic layer with microtubercles was secreted by the exterior epithelium after the completion of the main ammonitella wall by the interior shell gland. Thus, the early ammonite embryo might have had an endocochliate structural plan as in coleoids, and at the time of hatching the overlying mantle epithelium had shifted anteriorly to become an ectocochliate condition.     

腹足类个体发育中壳质结构的变化

腹足类个体发育中壳质结构的重要特点是壳质层的微观变化，包括原有壳质层的增厚、增生与上覆超微结构相同的壳质层、增生新的超微结构层、壳质层的相互消长与显微结构的演变。壳质层的微观变化决定了壳饰形成的４种类型：增厚型、刺顶型、刺穿型、叠覆型。叠覆的交错片体具加强贝壳抗破裂功能；交错片体排列方式的变化或新的显微结构层的出现均具分类意义。交错针状结构源于纤柱结构，纤柱结构又由简单柱状结构演变而来。     

Microstructural details in shells of the gastropod genera Carychiella and Carychium of the Middle Miocene

Microstructural details are revealed via scanning electron microscopy (SEM) in two carychiid species from the early Middle Miocene of Styria, SE Austria. The protoconchs of the shells of Carychiella eumicrum (Bourguignat 1857) and Carychium gibbum (Sandberger 1875) show different types of microstructure on the embryonic shell during ontogeny. Total, superficial punctate structure on the shell of Carychiella eumicrum contrasts with the protoconch–teleoconch demarcation (p/t boundary) observed on the protoconch of Carychium gibbum. Both species exhibit aragonitic microstructure. Diagenetic effects, prismatic, homogeneous and crossed lamellar microstructures are detectible in both species. Rheomorphic folding and dense pitting within the columella of Carychiella eumicrum suggest a structure–function relationship for tensile strength and bulk weight reduction in carychiid snails. We hypothesize that total superficial pitting on the shell of C. eumicum, seen here for the first time in the Carychiidae, suggests paedomorphosis as a life‐history strategy to palaeoecological conditions of the Rein Basin during the early Middle Miocene.     

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.     

Molecular phylogeny of protobranch bivalves and systematic implications of their shell microstructure

Higher systematics and evolutionary history of Protobranchia, a subclass of Bivalvia, have long been controversial due to paucity of prominent shell characters and difficulties in collecting live material for diverse taxa. Here, we evaluate the reliability of shell microstructure for protobranch higher systematics by reconstructing a molecular phylogeny of the subclass. Relationships were assessed using the nuclear (18S rRNA, 28S rRNA and histone H3) and mitochondrial (16S rRNA and cytochrome c oxidase subunit 1) gene sequences from 89 in-group species. Maximum likelihood reconstruction with the nuclear markers recognized five superfamilies (Nuculoidea, Solemyoidea, Manzanelloidea, Nuculanoidea and Sareptoidea) as the in-group clades of the monophyletic Protobranchia. Sareptoidea is herein redefined to comprise Sarepta and Setigloma in the sole family Sareptidae, whereas Pristigloma and its monotypic Pristiglomidae are transferred from this superfamily to Nuculanoidea, both in the order Nuculanida. Mapping of shell microstructure characters on the tree confirmed their conservativeness at superfamily level when only living species were taken into account. The Nuculoidea have shells with the outer prismatic and middle/inner nacreous structures; Solemyoidea are characterized by either the radially elongate simple prismatic structure or the reticulate structure in the outer shell layer; Manzanelloidea, Nuculanoidea and Sareptoidea have shells of homogeneous, fibrous prismatic and/or fine complex crossed lamellar structures, all of which lack large structural units. Our Bayesian time calibration, on the contrary, suggested frequent loss of nacre in the Paleozoic and Mesozoic history of Protobranchia, at least once each in Nuculoidea, Manzanelloidea, Solemyoidea and Sareptoidea in the Paleozoic, and perhaps multiple times in Nuculanoidea by the Mesozoic.     

Geometrical and crystallographic constraints determine the self-organization of shell microstructures in Unionidae (Bivalvia: Mollusca)

Unionid shells are characterized by an outer aragonitic prismatic layer and an inner nacreous layer. The prisms of the outer shell layer are composed of single-crystal fibres radiating from spheruliths. During prism development, fibres progressively recline to the growth front. There is competition between prisms, leading to the selection of bigger, evenly sized prisms. A new model explains this competition process between prisms, using fibres as elementary units of competition. Scanning electron microscopy and X-ray texture analysis show that, during prism growth, fibres become progressively orientated with their three crystallographic axes aligned, which results from geometric constraints and space limitations. Interestingly transition to the nacreous layer does not occur until a high degree of orientation of fibres is attained. There is no selection of crystal orientation in the nacreous layer and, as a result, the preferential orientation of crystals deteriorates. Deterioration of crystal orientation is most probably due to accumulation of errors as the epitaxial growth is suppressed by thick or continuous organic coats on some nacre crystals. In conclusion, the microstructural arrangement of the unionid shell is, to a large extent, self-organized with the main constraints being crystallographic and geometrical laws.     

Functional interpretation of inner shell layers in Triassic ceratid ammonites

The wrinkle layer the inner prismatic layer are described in three Triassic ceratid genera: Phyllocludiscites. Megaphyllites Proarcestes. Both layers have their counterparts in the shell wall of the recent Nautilus: the wrinkle layer corresponds to the mantle-adhesive layer the inner prismatic layer to the myostracal layer in Nautilus. A detailed structural functional comparison between these layers is given. The wrinkle layer is also compared with the oblique prismatic layer in recent gastropods.     

Description and quantification of pteropod shell dissolution: a sensitive bioindicator of ocean acidification

Anthropogenic ocean acidification is likely to have negative effects on marine calcifying organisms, such as shelled pteropods, by promoting dissolution of aragonite shells. Study of shell dissolution requires an accurate and sensitive method for assessing shell damage. Shell dissolution was induced through incubations in CO₂‐enriched seawater for 4 and 14 days. We describe a procedure that allows the level of dissolution to be assessed and classified into three main types: Type I with partial dissolution of the prismatic layer; Type II with exposure of underlying crossed‐lamellar layer, and Type III, where crossed‐lamellar layer shows signs of dissolution. Levels of dissolution showed a good correspondence to the incubation conditions, with the most severe damage found in specimens held for 14 days in undersaturated condition (Ω ~ 0.8). This methodology enables the response of small pelagic calcifiers to acidified conditions to be detected at an early stage, thus making pteropods a valuable bioindicator of future ocean acidification.     

Microstructure and mineralogy of the outer calcareous layer in the lower jaws of Cretaceous Tetragonitoidea and Desmoceratoidea (Ammonoidea)

Tanabe, K., Landman, N.H. & Kruta, I. 2011: Microstructure and mineralogy of the outer calcareous layer in the lower jaws of Cretaceous Tetragonitoidea and Desmoceratoidea (Ammonoidea). Lethaia, Vol. 45, pp. 191–199. Based on the differences in their relative size, overall shape, structure and the degree of development of an outer calcified covering, lower jaws of the Ammonoidea have been classified into four morphotypes: normal, anaptychus, aptychus and rhynchaptychus types. However, detailed microstructural and mineralogical comparison of these morphotypes has not yet been addressed. This article documents the results of SEM and XRD observations of the lower jaws of three Late Cretaceous ammonoid species belonging to the Tetragonitoidea (Anagaudryceras limatum) and Desmoceratoidea (Pachydiscus kamishakensis and Damesites aff. sugata), based on excellent material preserved in situ within the body chamber, and retaining an aragonitic shell wall. The lower jaws of the three species are assigned to an intermediate form between anaptychus and aptychus types for the first two species and the rhynchaptychus type for the third species. Their black, presumably originally chitinous outer lamella is wholly covered with a calcareous layer. The calcareous layer is composed of aragonite in D. aff. sugata and A. limatum, and calcite in P. kamishakensis. The microstructure of the outer calcareous layer differs among the three species, i.e. granular in A. limatum, spherulitic prismatic in D. sugata, and prismatic in P. kamishakensis, all of which can be distinguished from the lamellar and spongy structure of the outer‐paired calcitic plates of the typical aptychus‐type lower jaws in some Jurassic and Cretaceous Ammonitina and Ancyloceratina. Our study suggests that most Jurassic and Cretaceous ammonoids possessed an outer calcareous layer in their lower jaws, although its mineralogy, microstructure and relative thickness vary among different taxa. □Ammonoidea, Cretaceous, Desmoceratoidea, lower jaw, microstructure, Tetragonitoidea.     

Crystallographic characterization of the crossed lamellar structure in the bivalve Meretrix lamarckii using electron beam techniques

To understand the formation mechanism of crossed lamellar structures in molluskan shells, the crystallographic structural features in the shell of a bivalve, Meretrix lamarckii, were investigated using scanning electron microscopy, electron backscattered diffraction, and transmission electron microscopy with a focused ion beam sample preparation technique. Approximately 0.5 μm-thick lamellae (the second-order units) are piled up obliquely toward the growth direction to form the first-order unit and the obliquity is inverted between adjacent units along the shell thickness direction. The first-order units originate around the center of the shell, initially growing parallel to the shell and subsequently curving toward the inner or outer surfaces. The lamellae consist of aragonite granular and columnar layers, which group together to adopt the same crystal orientation forming crystallographic units (crystallites). Multiple {1 1 0} twins are common both in the granular and columnar layers. The crystallite c-axis is parallel to the columns and is inclined at angles 0–50° from the lamellar normal (dispersing among individual lamellae), toward the shell growth direction. Probably, the directions of the a- and b-axes are random in the lamellae, showing no specific orientation.     

Post-larval and larval shells ofJuranomia Fürsich & Werner 1989, andAnomia Linnaeus 1758 (Anomüdae, Bivalvia)

Post-larval and larval shells ofJuranomia calcibyssata from the Bathonian to Callovian of Poland and RecentAnomia membranacea from the Mediterranean are described and compared to other fossil and Recent members of the family Anomüdae. The stratigraphic range of the monospecific genusJuranomia, which, up to now, was only known from the Kimmeridgian, can be extended to include the Lower Bathonian. The state of preservation of the fossil species allows recognition of an internal aragonitic, branching and complex cross-lamellar shell layer in the post-larval left valve, which previous studies could only assume to be present.Anomia membranacea is a member of theA. ephippium lineage as proved, among other characters, by the presence of an outer calcitic prismatic layer in its right valve. It possesses an anterior pedal retractor in the left valve, which, in the original discussion of the phylogenetic affinities ofJuranomia, was thought to be lacking in species ofAnomia. Consequently, the generaJuranomia andAnomia only differ in two important shell characters: closeness or distance between the three central muscles and thickness of the inner aragonitic shell layer of the left valve. Larval shells ofJuranomia are similar to those of Recent anomiids in shape, size, the presence of a byssal notch in the right valve, and an external sinus and internal shelly process in the left valve. The last three features are parts of a single character which is considered as an autapomorphy of the stem species of the Anomiiudae. The small P I size ofJuranomia calcibyssata suggests a purely planktic-planktotrophic development and thus, high potential of dispersal, just as its modern counterparts. Irrespective of the general similarity in shell size, the mean dimensions of the P II are likely species-specific.     

Shell structure and affinity of vermiform 'gastropods'

Vermiform ‘gastropods’ are reported from a variety of rocks ranging from Givetian to Lower Triassic age. Examples encrusting shells and plants have been identified in non-marine shales, in addition to previously recognized occurrences in shallow marine microbial bioherms and stromatolites. SEM studies of the planorbiform or trochiform protoeonch reveal a shell wall comprising three calcite layers: an outer, initial acicular layer: a blocky prismatic layer: and an inner irregular micro-lamellar layer. Minor irregularities and microstructural details suggest a high original organic content. allowing flexibility for attachment. The sinistrally coiled (or hyperstrophic dextral) calcitie teleoconch is composed of an outer simple prismatic layer and inner micro-lamellar layer comprising sheets of irregular, platy, sometimes fused calcite tablets. displaying ridges and grooves similar to those of cross-bladed fabric. Repetition of layers may occur. Regular closely-spaced punctae. passing through and disrupting the micro-lamellar layer. are unlike any mollusean tubulation. Punetation may he a shell-strengthening response to uncoiling. Septa bear a centrai, anteriorly-projecting, probebly perforate protrusion. reminiscent of the siphunele of cephalopods and similar structures in tentaculitoida. The micro-lamellar layer in the protoconch, the micro-lamellar layer with distinctive ridge and groove structure and punetation in the teleoconch, and the structure of the septa point to a close affinity between vermiform ‘gastropods’ and the Tentaculitoidea. The three-layered microstructure of the protoconch and the coiled nature of the shell distinguish vermiform ‘gastropods’ from tentaculitoids. A consideration of the shared characters indicates that the tentaculitoids and vermiform ‘gastropods’ should be regarded as a sister group to the molluses. ***Vermiform gastropods: microstructure, protoconch, teleoconch, acicular laver, prismatic laver, micro-lameller layer. cross-bladed structure. punetation, septation. Tentaculitoidea     

Shell microstructure and mineralogy of the Littorinidae: ecological and evolutionary significance

An examination of the shell microstructure and mineralogy of species from 30 of the 32 genera and subgenera of the gastropod family Littorinidae shows that most species have a shell consisting of layers of aragonitic crossed-lamellar structure, with minor variations in some taxa. However, Pellilitorina, Risellopsis and most species of Littorina have partly or entirely calcitic shells. In Pellilitorina the shell is made entirely of calcitic crossed-foliated structure, while in the other two genera there is only an outer calcitic layer of irregular-prismatic structure. A cladistic analysis shows that the calcitic layers have been independently evolved in at least three clades. The calcite is found only in the outermost layers of the shell and in species inhabiting cooler waters of both northern and southern hemispheres. Calcium carbonate is more soluble in cold than warm water and, of the two polymorphs, calcite is about 35% less soluble than aragonite. We suggest that calcitic shell layers are an adaptation of high latitude littorinids to resist shell dissolution.     

New data on the enigmatic Ocruranus–Eohalobia group of Early Cambrian small skeletal fossils

Abstract: The Ocruranus–Eohalobia group, whose members were variously considered to be brachiopods, bivalves, chitons, tommotiids and coeloscleritophorans, are difficult to classify because of lack of morphological detail and evidence for skeletal reconstruction. New specimens from South China reveal more information about Ocruranus–Eohalobia and allow progress towards deciphering the skeletal reconstruction and phylogenetic affinity of this enigmatic group. Many specimens have a phosphatic inner and outer coat (mould) with empty space in between that resulted from dissolution of the original shell. Moreover, many of the internal moulds show a previously unknown type of shell microstructure that consisted of stacked layers of highly organized, acicular crystallites that radiated from the apex of the shell towards the aperture. The dissolved shell and needle‐like crystals suggest an original calcareous, probably aragonitic, shell mineralogy. A few specimens also show a polygonal texture in regions that suggests the shell had a thin, prismatic inner shell microstructure. Ocruranus and Eohalobia belong to the same skeleton, and we herein synonymize Eohalobia with the older Ocruranus. Moreover, new specimens from Meishucun reveal a third type of shell plate, similar in form and inferred placement to intermediate valves of chitons. Ocruranus is likely a mollusc, and possibly a member of the chiton stem lineage. If so, then the beginning of the known record of chitons would be extended back from late Cambrian (Saukia Zone; Furongian) to early Cambrian (Meishucunian; Series 1).     

Structure and composition of the aragonitic crossed lamellar layers in six species of Bivalvia and Gastropoda

The microstructures, the chemical composition and the soluble organic matrices of the aragonitic crossed lamellar layers of the shells of six species of molluscs have been studied. The microstructures and chemical contents are similar, whereas the quantities of organic matrices are variable. All the soluble matrices are glycoproteins, with low S contents. Their molecular weights, the protein-sugar ratios and acidities are variable. Neither a gastropod nor a bivalve pattern is recognized. The diversity of the organic matrices probably plays a main role in the fossilization processes of mollusc shells.