ORIGIN OF LAMINAR-SHELLED ARTICULATE BRACHIOPODS

The calcareous shell of Billingsella is composed of a primary layer of crystallites commonly disposed vertically or at high angles to the shell surface, and a laminar secondary layer of flat-lying blades which usually amalgamate laterally to form a succession of plates. The succession is unlike that of other billingsellaceans such as Nisusia or contemporaneous orthaceans like Oligomys and Orusia which have a secondary layer composed of fibres. It is, however, closely comparable with the impunctate, laminar secondary layer of the Triplesiacea and early Davidson-iacea. The other pene-contemporaneous laminar-shelled articulates, the Stropho-menacea and Plectambonitacea, differ in being pseudopunctate. These differences in shell structure suggest that the Strophomenida were polyphyletically derived, mainly from the Nisusiidae but also from the Billingsellidae which gave rise to the Davidsoniacea (and Triplesiacea).     

Shell structure and affinities of the enigmatic Lower Ordovician articulate brachiopod Lycophoria Lahusen, 1886

Wright, A. D. 1992 04 15: Shell structure and affinities of the enigmatic Lower Ordovician articulate brachiopod Lycophoria Lahusen. 1886. Lethaia . Vol. 25, pp. 125–129. Oslo. ISSN 0024–1164.
The enigmatic Baltoscandian Lower Ordovician brachiopod Lycophoriu has a combination of morphological characters that makes it difficult to place taxonomically. The more recent assignments of the genus have been with the Porambonitacea, the Triplesiacea and the Orthacea. A basic character in articulate brachiopods is the differentiation of the secondary shell into either stacked fibres or laminar sheets. The hitherto unknown shell structure in Lycophoria has been examined under the electron microscope and is shown to be fibrous, which is taken as ruling out any close affinity with the lamellar shelled Triplesiacea. Despite superficial similarities, features of the shell interior are not compatible with the pentameride Porambonitacea and although there are differences from the typical orthacean, these are no greater than those of the accepted orthid Producrorrhis. Lycophoria , in the monotypic family Lycophoriidae, is accordingly best regarded as a specialized offshoot of the basic orthacean stock. * Shell microstructure, Lycophoridae, Orrhacea, Porambonitacea, Triplesiacea, Lower Ordovician, Baltoscandia .     

Tentaculitoid affinities of the tubeworm-like fossil Tymbochoos sinclairi (Okulitch, 1937) from the Ordovician of North America

Tymbochoos sinclairi (Okulitch, 1937) has a laminar tube structure and pseudopuncta similar to the tentaculitoids. This suggests that Tymbochoos belongs to the Tentaculitoidea Bou?ek, 1964.     

Microborings in Recent brachiopods and the functions of caeca

Microborings in the primary shell layer of Recent brachiopods are clearly seen to avoid endopunctamicroscopic canals pervading the shell fabric and housing papillose extcnlions of the mantle (the caeca). This avoidance confirms the suggestion that the caecal contents inhibit boring organisms (Owen & Williams 1969; Proc. R. Soc. Loud. B, 172 ), and as such the caecum can be considered as an important instrument in protecting the brachiopod shell. A comparison of the relative fecundity of co-habitating impunctate and cndopunctate New Zealand brachiopods provides indirect evidence that the caecum may indeed also function in a nutrient storage capacity. Brachiopods, microborings, primary shell layer, endopuncta, defence, storage.     

Evolutionary and Diagenetic Changes in the Chemico-structure of the Shell of Cranioid Brachiopods

The laminar dorsal valve of living Neocrania consists of: a primary layer of rhombohedral tablets, composed of granular calcite and commonly forming slats orthogonal to the margin, associated with polysaccharides and a fibrous 60 kDa protein; and a secondary layer of spirally growing (10.4) rhombohedra, doped with the 60 kDa protein and interleaved with membranes of a fibrous 44 kDa protein. The ventral valve consists exclusively of a primary layer with the same composition and basic structure as that of the dorsal valve. Investigation of selected antecedents shows that the chemico-structure of the Neocrania shell has been virtually unchanged since the first appearance of the stock in the Early Ordovician (Arenig). The greatest phylogenetic change affected the ventral valve that varied, even in Ordovician genera, from a film of calcitic blades to a complete succession of primary and secondary laminae. The most profound diagenetic changes occurred before the Late Cretaceous with proteins degrading into peptides that were dispersed, with the loss of less stable amino acids, during laminar recrystallization. Palaeozoic shells suffered further recrystallization but, even after pressure solution, the original laminar fabric was replicated long after it had lost its constraining organic membranes.     

Description of the early ontogenic part of the Tentaculitids, with implications for classification

Ontogenic development and classification of tentaculitids at high systematic levels are reevaluated in the light of new findings on shell structure and morphology of larval parts, and these features are here regarded as being of primary importance for taxonomy. Class Tentaculita Bouček, 1964 is subdivided into two subclasses, of which subclass Chionioconarida Farsan, 1994 is distinguished by a tubular larval process closed at the apex and covered with microrings. The process is differentiated into a prolarval, metalarval and epilarval part, of which the latter coincides with metamorphosis. Morphology of the larval parts suggests that metamorphosis proceeds in two different manners, giving rise to superorders within this subclass. Within superorder Trompetoconarida Farsan, 1994 a bilaterally symmetrical larval cone develops with an aperture oblique to the long axis of the conch; following metamorphosis the conch becomes radially symmetrical and the aperture perpendicular to the axis; secondary shell, septa and pseudopunctae develop in the adult phase, and the structure of the shell is lamellar. In contrast, within the second superorder, Lirioconarida Farsan, 1994 the epilarval tube develops into a larval bulb with no changes in symmetry and position of the aperture; secondary shell, septa and pseudopuncta are absent. The microstructure of the shell is lamellar in the larval part whereas in postlarval parts it is either sigmoidal or lamellar. The subclass Dacryoconarida Fisher, 1962 possesses a subspherical, tear- or drop-like embryonic chamber which may have a caudal process. The microstructure of the embryonic chamber is variable within this group, being lamellar in some taxa whereas in others, a single layer of shell is present. The postembryonic parts of the lamellar forms possess nacreous or sigmoidal structures.     

Microstructure of the linguliformean brachiopod Linnarssonia from the Lower Cambrian of Sichuan, China

Acrotretoids, one of the oldest brachiopod groups, are abundant in the Lower Cambrian Jiulaodong Formation. The shell of Linnarssonia sp. is composed of two layers: a primary layer and a columnar secondary layer. The primary layer is mostly exfoliated, resulting in exposure of the openings to the central canal of the columns. Filae are seen on the surface of the columnar layer, indicating that the columnar secondary layer has influenced changes in ornament on the shell surface. The larval shell has only very weak ripples; the post-larval shell has obvious concentric ribs. Small pits of variable shape cover almost the entire shell surface. The secondary layer is composed of several columnar laminations, each of which comprises both the upper and lower laminae and the cylindrical columns between them. On the inner side of shell the thin columnar laminations increase. The new microstructural data show that two shell layers are developed in Early Cambrian acrotretoid brachiopods; the columnar lamination may be a primitive feature of the microstructural development of the Brachiopoda and may help establish the affinity between different stem-group brachiopods.     

A scanning electron microscopic study of the inorganic and organic matrices comprising the mature shell of Amblema, a fresh-water mollusc

The scanning electron microscope has been used to describe the morphology of the mature shell in a fresh-water bivalve. The structure of the organic and inorganic components within the nacre, the myostracum, and the prismatic layer is described. A transitional or intermediate zone, interposed between the prismatic layer and the nacre, was identified. In demineralized samples, the organic component of the nacre was found to consist of parallel matricial sheets interconnected by irregular transverse bridges. The structure of the mineral component of the nacre was found to vary with the method of specimen preparation. With polished-etched samples, brick-like units were seen. When shells were simply broken and fixed in osmium, the layers of nacreous material consisted of fusing rhomboidal crystals of aragonite which demonstrated subconchoidal fractures. On the inner surface of the shell, the rhomboidal crystals showed an apparent spiral growth pattern. The myostracum was characterized by regions of modified nacreous structure consisting of enlarged aragonite crystals with a pyramidal morphology. The peripheral aspect of the muscle scars was characterized by rhomboidal crystals, the latter fusing to form the typical nacreous laminae. The uniqueness of the anterior adductor scar is exemplified by the presence of pores, each pore walled by pyramidal units, for the insertion of adductor fibres. In most regions of the shell, the prismatic layer consisted of one prism unit thickness with a height of approximately 225–250 μm. However, in two specialized regions of the shell, this layer was seen to consist of multiple layers of stacked prisms. The organic matrices of the prismatic layer are arranged in a honeycomb-like arrangement and packed with mineralized spherical subunits.     

Shell structure of the Early Cretaceous (Berriasian-Barremian) Rhynchonellids from Dagestan

The shell structure of five rhynchonellid species from the Lower Cretaceous of Dagestan was studied for the first time. The primary layer is rarely preserved in fossil rhynchonellids. Most of the species studied have a primary layer composed of prismatic calcite, one species has a finely granulated primary layer. The secondary fibrous layer consists of crossing bundles of fibers the (throughout shell thickness) and parallel bundles of fibers on the valve bottom.     

A HISTORY OF SKELETAL SECRETION AMONG ARTICULATE BRACHIOPODS

Studies of the ultrastructure of the exoskeleton of Notosaria nigricans (Sowerby), which can be used as the standard succession for articulate brachiopods, show that shell secretion involves six distinct operations giving rise to the following layers: mucopolysaccharide, outer fibrillar triple-layered membrane, mucoprotein, inner fibrillar triple-layered membrane, calcareous primary layer and calcareous-organic secondary layer. Comparison with the secretory régimes of terebratulids like Waltonia inconspicua (Sowerby) suggests that only four of these operations are fundamental, those controlling secretion of mucopolysaccharide, outer fibrillar membrane and the primary and secondary shell. An endoskeletal secretory phase giving rise to spicules in the mantle and lophophore is also found in many terebratulids. In the geological record, the orthodox primary-secondary shell can be traced back to the billingsellaceans and is presumed to have been associated with the mucopolysaccharide layer and a fibrillar membrane which together might well have constituted the prototypic organic exoskeleton. Deviation from this pattern during brachiopod evolution was relatively minor and was, so far as is known, limited to the strophomenids, some spiriferids and thecideidines.     

The “rostrum”-problem in coleoid terminology – an attempt to clarify inconsistencies

In the coleoid literature, the terminology of shell elements that are deposited on the external surface of the primary shell wall is inconsistent and confusing. Morpho- and phylogenetic interpretations have been therefore controversial. A strict layer-by-layer comparison of seventeen species from seven coleoid subgroups suggests that the coleoid shell is covered by either one (Diplobelida, Spirulida, Sepiida, Vampyropoda) or two (Aulacocerida, Belemnitida, Belemnoteuthidida) outer shell formations. The confusion has been caused mainly by the often ignored presence of a primordial rostrum in the Aulacocerida, Belemnitida, and Belemnoteuthidida. The primordial rostrum is a secondary shell formation, which covers the entire primary shell and, which is itself enveloped by a tertiary shell formation, the rostrum proper. In the Diplobelida, Spirulida, and Sepiida, the primary shell is invested by a single outer formation, the sheath; a rostrum proper is absent in the latter groups. As a secondary shell formation, the aulacocerid, belemnitid and belemnoteuthidid primordial rostrum on the one hand, and the diplobelid, spirulid and sepiid sheath on the other hand are considered to represent homologues. Accordingly, the rostrum proper of the Aulacocerida, Belemnitida, and Belemnoteuthidida are homologues as tertiary shell formations. Outer shell formations in gladius-bearing vampyropods (and teuthids?) might be represented by a single layer. The clarification of the homology of secondary and tertiary shell formations, however, cannot resolve phylogenetic relationships within the Coleoidea.     

THE MANTLE AND SHELL OF SOLEMYA PARKINSONI (PROTOBRANCHIA: BIVALVIA)

The shell of Solemya exhibits considerable flexibility which is further enhanced by the marked extension of the periostracum beyond the calcareous portions of the valves. This fcature, more than any other, has made possible the habit, unique among bivalves, of burrowing deep within the substrate without direct contact with the water above. The inner calcareous layer of tho valves is restricted to a small area near the umbones while the outer calcareous layer is thin and contains a high proportion of organic material. The shell conchiolin consists mainly of protein, varying in composition, but much of it strengthcned by quinone-tanning, and in ccrtain regions probably by the presence of appreciable quantities of chitin. The ligament, although superficially resembling an amphidetic structure, is opisthodetic, the extcnsion anterior to the umbones consisting of anterior outer layer only.
The mantle is characterized by an extension of the outer fold of the mantle margin which has effected equally both the inner and outer surfaces of this fold. The secretory epithelium and the modified pallial musculature, contraction of which results in the intucking and plaiting of the periostracum, is dcscribed. Simple tubular oil glands open at the mantlo margin and are responsible for the water-repellent nature of the periostracum.
The form of the mantlelshell and that of the enclosed body are discussed and compared with those of other bivalves in which elongation of the mantle/shell is achieved in a different way. It is concluded that the mantlelshell of Solemya is of little value in determining its relationships, and that the greatly elongatod ligament, the edentulous hinge and the flexible shell are all adaptations to a specialized mode of life.     

Shell microstructure of <Emphasis Type="Italic">Discinisca suborbicularis</Emphasis> sp. nov. (Brachiopoda,Lingulata) from the Upper Jurassic of Western Siberia

A new species of the phosphatic brachiopod Discinisca suborbicularis (family Discinidae) from the Upper Jurassic of Western Siberia (latitudinal Priob’e) is established. The shell microstructure under protegulum and brephic shell and the microstructure of adult shell (that corresponds to three developmental stages) are described in detail. The shell microstructure of new species considerably differs from Paleozoic discinids and is similar to Recent discinids. D. suborbicularis differs from recent discinids in the presence of protegulum and thicker organophosphatic primary layer.     

淡水贝类贝壳多层构造形成研究

对几种淡水贝（包括蚌、螺）进行形态及组织学观察，并通过实验方法重现贝壳三种物质，即：角质、棱柱质、珍珠质的生成过程，结果表明：外套膜外表皮细胞是由相同类型细胞组成，这些相同细胞在不同的作用条件下形成贝壳多层构造。     

Muscle formation in the sexual genetation of Intoshia variabili (Orthonectida)

Muscle formation in Intoshia variabili (Orthonectida) has been studied in the course of development of the sexual (free-living) specimen. The muscle system originates in the early embryogenesis as a distinct continuous layer located between the outer cell layer and the inner cell mass. Later this cell layer disintegrates into separate muscle strips. The presence of a distinct muscle system in Orthonectida and the pattern of its development evidences for placing this group into Triploblastica rather than into Diploblastica.     

The ontogeny of shell secretion in Terebratalia transversa (Brachiopoda, Articulata). II. Formation of the protegulum and juvenile shell

The fine structure of the shell and underlying mantle in young juveniles of the articulate brachiopod Terebratalia transversa has been examined by electron microscopy. The first shell produced by the mantle consists of a nonhinged protegulum that lacks concentric growth lines. The protegulum is secreted within a day after larval metamorphosis and typically measures 140-150 micron long. A thin organic periostracum constitutes the outer layer of the protegulum, and finely granular shell material occurs beneath the periostracum. Protegula resist digestion in sodium hypochlorite and are refractory to sectioning, suggesting that the subperiostracal portion of the primordial shell is mineralized. The juvenile shell at 4 days postmetamorphosis possesses incomplete sockets and rudimentary teeth that consist of nonfibrous material. The secondary layer occuring in the inner part of the juvenile shell contains imbricated fibers, whereas the outer portion of the shell comprises a bipartite periostracum and an underlying primary layer of nonfibrous shell. Deposition of the periostracum takes place within a slot that is situated between the so-called lobate and vesicular cells of the outer mantle lobe. Vesicular cells deposit the basal layer of the periostracum, while lobate cells contribute materials to the overlying periostracal superstructure. Cells with numerous tonofibrils and hemidesmosomes differentiate in the outer mantle epithelium at sites of muscle attachments, and unbranched punctae that surround mantle caeca develop throughout the subperiostracal portion of the shell. Three weeks after metamorphosis, the juvenile shell averages about 320 micron in length and is similar in ultrastructure to the shells secreted by adult articulates.     

The problematic cemented Devonian brachiopod Schuchertellopsis durbutensis Maillieux, 1939

The Devonian cemented brachiopod Schuchertellopsis durbutensis has proved difficult to classify and its possible taxonomic relationships are unknown. Morphologically Schuchertellopsis resembles more closely members of the Orthotetidina than the Davidsoniidina. Examination of the shell structure, a key diagnostic feature of the Orthotetidina, shows that Schuchertellopsis has the cross laminar secondary shell typical of all orthotetidines. However, the presence of both pseudopunctate and incipient an extropunctate fabric within the ventral valve is unique amongst orthotetidine brachiopods and is thought to represent a phase of shell fabric experimentation. Schuchertellopsis probably fits most comfortably within the Schuchertellidae, and is the earliest representative of that family.     

四川中侏罗世的龟化石

本文对产自四川自贡中侏罗世的三件龟甲标本作了鉴定,认为均可归成渝龟属(Chengyuchelys)。一为自贡成渝龟,新种(Chengyuchelys zigongensis,sp.nov.),一为似贝氏成渝龟(C.baenoides, Young et Chow),一为成渝龟未定种(Chengyuchelys sp.)。在形态描述的基础上,讨论了成渝龟属的系统分类。     

Skin structure and cornification proteins in the soft-shelled turtle Trionyx spiniferus

In contrast to most chelonians, the fully aquatic soft-shelled turtles have a smooth, unscaled, and pliable shell. The skin of the shell, tail, limbs, and neck of juveniles of Trionyx spiniferus has been studied by ultrastructural, immunocytochemical, and immunoblotting methods. The epidermis of the carapace and plastron has a thick corneous layer composed of alpha-corneocytes surrounded by a cornified cell envelope. The softer epidermis is similar to that of the shell but the epidermis and corneous layer are much thinner. Pre-corneous cells in both soft and shell epidermis are rich in vesicles produced in the Golgi apparatus and smooth endoplasmic vesicles, and contain numerous dense-core mucus-like and vesicular (lamellar) bodies. Secreted material is present among corneocytes where it probably forms an extensive intercellular lipid-mucus waterproof barrier. The dermis is very thick and composed of several layers of collagen bundles that form a plywood-patterned dermis. This dermis constitutes a strong mechanical barrier that compensates for the low content in beta-keratin, and lack of cornified scutes and dermal bones. The growth of the shell mainly occurs along the lateral margins. Immunocytochemistry reveals the presence of some beta-keratin in soft and shell epidermis, and this is confirmed by immunoblotting where bands at 18 and 32-35 kDa are present. Other proteins of the cornified cell envelope (loricrin and sciellin) or associated to lipid trafficking (caveolin-1) are also detected by immunoblotting. Loricrin positive bands at 24 and 57 kDa are present while bands cross-reactive for sciellin are seen at 45 and 53 kDa. Caveolin-1 positive bands are seen at 21-22 kDa. The presence of these proteins indicates that the epidermis is both coriaceous and waterproof. These results suggest that the shell of Trionyx is tough enough to be as mechanically efficient as the hard shell of the other turtles. At the same time, a soft shell is lighter, its shape is more easily controlled by muscles, and it allows a more controlled closure and retraction of limbs and neck inside the shell. Thus, the shell makes the animal more streamlined for swimming.