The relationship of the mantle and shell of the Polyplacophora in comparison with that of other Mollusca

The structure and growth of the polyplacophoran shell, characteristically consisting of eight plates surrounded by a girdle, is examined in the light of current views on the relationships of mantle and shell in the Bivalvia. The periostracum and outer and inner calcareous layers of the shell of the latter group are homologous with the cuticle, tegmentum and articulamentum respectively of the shell of the Polyplacophora. The margin of the mantle consists of a large marginal fold, which secretes the cuticular girdle, and a small accessory fold bearing mucous cells. These are functionally comparable with all three folds of the mantle margin found in other molluscs, although anatomically the marginal fold of the chitons probably represents only the inner surface of the outer fold of the mantle margin.
The cuticle not only forms the girdle, which bears calcified spines or spicules, but also extends between the shell plates. The principal part of the cuticle consists largely of mucopolysaccharide material but there is also a thin discrete inner region which is similar chemically to the periostracum of other molluscs. The cuticle, possibly without spines, probably covered the entire dorsal surface of a primitive placophoran and beneath this, plates developed. As these grew the cuticle became worn away except marginally and between the plates. It is suggested that a covering of mucus over the visceropallium may have been the forerunner of the molluscan shell and the possible evolutionary relationships of the shell throughout the Mollusca are discussed.     

The relationship of the mantle and shell of the Polyplacophora in comparison with that of other Mollusca

The structure and growth of the polyplacophoran shell, characteristically consisting of eight plates surrounded by a girdle, is examined in the light of current views on the relationships of mantle and shell in the Bivalvia. The periostracum and outer and inner calcareous layers of the shell of the latter group are homologous with the cuticle, tegmentum and articulamentum respectively of the shell of the Polyplacophora. The margin of the mantle consists of a large marginal fold, which secretes the cuticular girdle, and a small accessory fold bearing mucous cells. These are functionally comparable with all three folds of the mantle margin found in other molluscs, although anatomically the marginal fold of the chitons probably represents only the inner surface of the outer fold of the mantle margin.
The cuticle not only forms the girdle, which bears calcified spines or spicules, but also extends between the shell plates. The principal part of the cuticle consists largely of mucopolysaccharide material but there is also a thin discrete inner region which is similar chemically to the periostracum of other molluscs. The cuticle, possibly without spines, probably covered the entire dorsal surface of a primitive placophoran and beneath this, plates developed. As these grew the cuticle became worn away except marginally and between the plates. It is suggested that a covering of mucus over the visceropallium may have been the forerunner of the molluscan shell and the possibleevolutionary relationships of the shell throughout the Mollusca are discussed.     

Periostracal mineralization in the gastrochaenid bivalve Spengleria

We investigated the spikes on the outer shell surface of the endolithic gastrochaenid bivalve genus Spengleria with a view to understand the mechanism by which they form and evaluate their homology with spikes in other heterodont and palaeoheterodont bivalves. We discovered that spike formation varied in mechanism between different parts of the valve. In the posterior region, spikes form within the translucent layer of the periostracum but separated from the calcareous part of the shell. By contrast those spikes in the anterior and ventral region, despite also forming within the translucent periostracal layer, become incorporated into the outer shell layer. Spikes in the posterior area of Spengleria mytiloides form only on the outer surface of the periostracum and are therefore, not encased in periostracal material. Despite differences in construction between these gastrochaenid spikes and those of other heterodont and palaeoheterodont bivalves, all involve calcification of the inner translucent periostracal layer which may indicate a deeper homology.     

Geometric and functional constraints on bivalve shell morphology

The range of shell morphologies available to bivalves is constrained by the geometric properties of coiled shells, and by two contrasting functional necessities: positioning the umbones at a distance from each other, to allow an adequate amount of shell gape, and limiting linear growth of the axial shell margin, in order to prevent the ligament from being rapidly stretched beyond its elasticity limits. These necessities are achieved, or circumvented, in one or more of the following ways: (1) evolving a range of inequivalve adaptations, (2) allowing a large amount of interumbonal growth, while simultaneously adopting a ligament that quickly breaks and is continuously replaced during ontogeny, (3) adopting an outward curving ligament which flexes along its entire width, thus effectively placing the actual pivoting axis of the valves dorsally to the axial shell margin, (4) substituting the ligament with diductor muscles, (5) stopping growth before shell coiling reaches half whorl, (6) decreasing the whorl expansion rate throughout ontogeny, (7) orienting the coiling axes of the umbones at an angle to each other and to the hinge axis.     

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.     

Differentiation and Growth of the Brachiopod Mantle

Cell differentiation in the mantle edge of Notosaria, Thecidelhnaand Glottidia, representing respectively, the impunctate andpunctate calcareous articulate and chitinophosphatic inarticulatebrachiopods, is described. Comparison of electron micrographssuggests that outer epithelium which secretes periostracum andmineral shell, is separated from inner epithelium by a bandof "lobate" cells, of variable width, exuding an impersistentmucopolysaccharide film or pellicle. The lobate cells alwaysoccupy the same relative position on the inner surface of theouter mantle lobe; but the outer epithelium is commonly connectedwith the inner surface of the periostracum by papillae and protoplasmicstrands which persist during mineral deposition and ensure thatboth shell and attached mantle remain in situ relative to theoutwardly expanding inner surface of the outer mantle lobe.In the prototypic brachiopod, the lobate cells are likely atfirst to have occupied the hinge of the mantel fold but laterto have been displaced into their present position by the rigidoutward growing edge of the mineral shell.     

Biology and functional morphology of a new species of endolithic Bryopa (Bivalvia: Anomalodesmata: Clavagelloidea) from Japan and a comparison with fossil species of Stirpulina and other Clavagellidae

Abstract. A new species of Clavagellidae, Bryopa aligamenta, from Okinawa, Japan, is described. The species is endolithic in living corals, with the left valve cemented to the crypt wall, as in all clavagellids. The free right valve exhibits an unusual growth pattern, with commarginal lines seemingly arising from the posterior valve margin and extending towards the anterior. This results from: (i) progressive anterior erosion of the umbones, probably as a consequence of the boring process; (ii) the apparent migration posteriorly, as the umbones are eroded, of the dorso‐ventral growth axis of the shell; and (iii) enhanced posterior inter‐commarginal growth. Unlike other clavagellid genera and species, however, there is no discernible primary ligament, at least in the adult. It is possible, however, that if a juvenile ligament were present (as in B. lata), it too would be lost as a consequence of antero‐dorsal erosion during boring. To retain valve alignment in the absence of a primary ligament, and possibly upon reaching an adult size, the mantle lays down alternating layers of calcium carbonate and proteinaceous periostracum onto the interior surface of the shell to thicken it, most noticeably marginally and, especially, posteriorly. The two valves are united dorsally, therefore, by thin layers of periostracum that probably exert a minimal opening force. B. aligamenta is, however, further characterised by large adductor, pallial, and siphonal retractor muscles so that the entire animal is encased tightly within an internally strengthened shell within a crypt. Movement must be minimal, blood being pumped into pallial haemocoels to push open the valves and extend the siphons. Despite a suggestion to the contrary, Bryopa is retained in the Clavagellidae, its unusual growth processes resulting from an endolithic life style within living corals. The fossil clavagellid Stirpulina bacillus, from the Pliocene/Pleistocene of Palermo, Sicily, Italy, was, unlike Bryopa aligamenta and other clavagellids, endobenthic, with a long adventitious tube and anterior watering pot superficially similar to species of Penicillidae, another family of the Clavagelloidea. Furthermore, as in all clavagellids only the left valve is fused into the fabric of the tube, the right being free within it. In all penicillids, both valves are fused into the fabric of their tubes. The watering pots of the fossil S. coronata, S. vicentina, and S. bacillus, moreover, are formed in a different manner to that of penicillids, by progressive encasement of the right valve inside the tube. In penicillids, the tube is secreted in a single event from the general mantle surface and the incorporation of both valves into its fabric. The constituent genera of the Clavagellidae thus constitute an example of parallel evolution with members of the Penicillidae.     

Ultrastructural studies of the mantle and the periostracal lamina in the manila clam, Ruditapes philippinarum

The four folds of the mantle and the periostracal lamina of R. philippinarum were studied using light, transmission and scanning electron microscopy to determine the histochemical and ultrastructural relationship existing between the mantle and the shell edge. The different cells lining the four folds, and in particular those of the periostracal groove, are described in relation to their secretions. The initial pellicle of the periostracum arises in the intercellular space between the basal cell and the first intermediate cell. In front of the third cell of the inner surface of the outer fold, the periostracal lamina is composed of two major layers; an outer electron-dense layer or periostracum and an inner electron-lucent fibrous layer or fibrous matrix. The role and the fate of these two layers differ; the outer layer will recover the external surface of the shell and the inner layer will contribute to shell growth.     

The biology and functional morphology of Pteria brevialata (Bivalvia: Pterioidea), epizoic on gorgonians in Hong Kong

Pteria brevialata characteristically attaches to Hicksonella princeps (Cnidaria: Gorgonoidea) at depths > 10 m in Hong Kong and the South China Sea. Attachment is by a stout byssus to the basal regions of the gorgonian colonies. The shell appears typical of the Pteriidae, but closer inspection shows that the antero-ventral margin, in particular, moulds itself to the form of the cylindrical gorgonian, which also changes its growth form, so that securer attachment is achieved. The margin of the right valve is not nacred so that the flexible outer prismatic layer adpresses firmly against the more extensively nacred left valve when they close; mantle and ctenidia also being withdrawn deeply. This is probably an anti-predation device. There are no mantle fusions, even between posterior inhalant and exhalant streams. The mantle margin is, however, complexly tentaculate. Pteria brevialata is monomyarian with reduced anterior byssal retractor muscles. The anterior face of the shell is thus greatly reduced whereas the posterior is inflated and postero-dorsally 'winged'.
Other features of the anatomy are described, including the ctenidial eyespot and simple photoreceptors in the inner component of the duplicated outer mantle folds, under the periostracum. Similar structures in other pterioideans and arcoideans suggest a close relationship between their respective orders.
It is concluded that gorgonian-associated pterioids can be derived from an epibyssate, crevice-dwelling ancestor, exploiting, as solitary individuals, the currents and clean oceanic waters above the substratum. This has three important advantages: (a) removal from surface-roving predators (the gorgonian providing additional protection); (b) exploitation of a niche hitherto unoccupied by bivalves and (c) removal of the animal from the sea-bed and exclusive exploitation of potential food held in suspension. Current stresses to attachment are avoided by modifications to the shell.     

The status of the Plicatulidae and the Dimyidae in relation to the superfamily Pectinacea (Mollusca: Bivalvia)

In the families Pectinidae and Propeamussidae (scallops) the massive rounded inner ligament layer which provides the opening thrust is bounded by long stretches of anterior and posterior outer ligament layer which, in the absence of teeth, maintain alignment of the valves. In the Spondylidae these outer layers migrate centrally to unite on either side of the inner layer, the primary ligament becoming transversely instead of longitudinally disposed. A secondary ligament of fused periostracum with no tensile strength unites the still long hinge line but secondary ball and socket teeth assume responsibility for alignment of the valves. In the Plicatulidae and the Dimyidae there is the same transverse disposition of the primary ligament but the mantle margins overarch this with production of a continuous longitudinal "external" secondary ligament above the "internal" primary ligament. Although resemblances with the Spondylidae are greater than with the Pectinidae and Propeamussidae, differences from all three are great enough to justify separation of the other two families into a new superfamily Plicatulacea. The monomyarian condition in the Plicatulidae could well have arisen from conditions similar to those in the Dimyidae, that is following cementation when dimyarian and not following byssal attachment as it does in the three families of the now proposed restricted Pectinacea.     

HYDRAULIC BURROWING IN THE BIVALVE MYA ARENARIA LINNAEUS (MYOIDEA) AND ASSOCIATED LIGAMENTAL ADAPTATIONS

The burrowing behaviour of the bivalve Mya arenaria from tidalflats of the Dutch Wadden Sea has been observed and recorded.Compared to other bivalves, M. arenaria is a very slow burrower,its burrowing behaviour being unique among bivalves since itis based essentially on the ejection of water through the pedalgape, with little assistance by the foot, which performs onlyan anchoring function. Water ejection is specially powerfuland individual jets may last several seconds, thus constitutingan effective means of removing sand from below the animal duringdigging. This hydraulic burrowing is more effective in loosesandy than in cohesive muddy substrates. Water ejection is providedby the ability of the bivalve to rock its valves across a dorsoventralaxis. This rocking motion implies special modifications of theligamental area. The ligament is conical in appearance and runsdorso-ventrally between the two chondrophores which are placedin two planes parallel to the cardinal axis. During rockingthe whole ligament acts in torsion and the lamellar layer ofthe ligament opposes closing of the anterior part of the valves.During normal adduction of the valves the ligament acts in bending,the axis of motion being placed internally with respect to thecardinal axis. This leads to approaching of the umbones withcomplete adduction and to resorption of the left umbo. Fromthe adaptive point of view, the slow hydraulic mode of burrowingis sufficient to cope with the slow sedimentation and erosionrates of the tidal flats in which M. arenaria lives. This burrowingmode implies the existence of a tiny foot, which leaves roomfor other organs within the mantle cavity. This, together withanterior divarication of the valves permits a large volume ofwater to be ejected from the mantle cavity, but, in the caseof M. arenaria, also the existence of an enormous stomach, possiblyas an adaptation for food processing. (Received 12 April 1996; accepted 2 October 1996)     

THE ANATOMY OF CALLOCARDIA HUNGERFORDI (BIVALVIA: VENERIDAE) AND THE ORIGIN OF ITS SHELL CAMOUFLAGE

Callocardia hungerfordi (Veneridae: Pitarinae) lives in subtidalmuds (220 to 240m C.D.) and is covered by a dense mat of mudthat, effectively, camouflages the shell. The periostracum is two layered. The inner layer is thick andpleated, the outer thin and perforated. From the outer surfaceof the inner layer develop numerous, delicate (0.5 mm in diameter),calcified, periostracal needles. These penetrate the outer periostracum.Mucus produced from sub-epithelial glands in the inner surfaceof the mantle, slides over the cuticle-covered epithelium ofthe inner and outer surfaces of the inner fold and the innersurface of the middle mantle fold to coat the outer surfaceof the periostracum and its calcified needles. Increased productionat some times produces solidified strands of mucus which bindmud and detrital material into their fabric to create the shellcamouflage. Calcified periostracal needles have been identified in othervenerids, including some members of the Pitarinae, but how theyare secreted and how the covering they attract is producedand, thus, how the whole structure functions, has not been explained. (Received 7 December 1998; accepted 5 February 1999)     

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.     

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。     

Enigmatic septa in shells of some Middle Jurassic Pholadomya (Bivalvia) from Poland

The interior of 36 specimens of Pholadomya Sowerby (Bivalvia) from the Middle Jurassic of Poland reveals the presence of unusual septa that separate sediment‐filled chambers from the shell interior. The septa occupy one or several recurrent loci in shells of various individuals, that is within umbones, in pallial sinuses and along the shell margins. Based on the location and shape of the septa, eight forms grouped into types and varieties are identified. A possible cause for the formation of septa is sediment toxicity, but intrusion of sediment to the shell interior must have been linked to shell breakage or rupture of the free periostracum. One form of septa occurs in perforated umbones, common in Pholadomya; other forms occur in intact shells, which suggest damage to free periostracum. The most likely cause for the latter is the presence of parasites, especially digenean trematodes, for which clams were intermediate hosts. The morphological aberrations presented here were hitherto unknown in both fossil and extant bivalves. This study is also the first report of pathologies in Anomalodesmata.     

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.     

Embryonic development of the shell in biomphalaria glabrata (Say).

During embryogenesis of the fresh water snail Biomphalaria glabrata (Say) (Pulmonata, Basommatophora) shell formation has been studied by light and electron microscopical techniques. The shell field invagination (SFI), the secretion of the first shell layers, the development of the shell-forming mantle edge gland and spindle formation have been investigated. During embryonic development at 28 degrees C environmental temperature, the shell field invaginates after 35 h. After 40 h the SFI is closed apically by cellular protrusions and scale-like precursors of the periostracum. The first electron translucent layer of the periostracum stems from electron dense vesicles of the cells which lie at the opening of the SFI. A second electron dense layer appears some hours afterwards. When the shell appears birefringent in the polarizing microscope (45 h of development) calcium can be detected in it using energy dispersive x-ray analysis. As calcification occurs the intercrystalline matrix appears under the periostracum and the SFI begins to open. In embryos of 60 h the mantle cavity appears at the left caudal side. When the mantle edge groove develops (65 h of development) lamellate units are added to the outer layer of the periostracum, but no distinct lamellar layer is formed in B. glabrata. In addition to the lamellar cell and the periostracum cell, a secretory cell can be observed in the developing groove. After 65 h of development, spindle formation starts and the shell begins to coil in a left hand spiral. After 5 days of development the embryos are ready to leave the egg capsules.     

REMOTE BIOMINERALIZATION IN DIVARICATE RIBS OF STRIGILLA AND SOLECURTUS (TELLINOIDEA: BIVALVIA)

Deposits composed of aragonite prisms, which were formed afterthe outer shell layer, have been found at the posterior steepslopes of divaricate ribs in two species of Strigilla and anothertwo of Solecurtus. These prisms have their axes oriented perpendicularto the outer shell surface and differ in morphology from fibresof the surface-parallel composite prisms forming the outer shell.They display crystalline features indicating that, unlike crystalsforming the outer shell surface, their growth front was free,unconstrained by the mantle or periostracum. These particulardeposits are called free-growing prisms (FGPs). In these generathe periostracum is clearly not the substrate for biomineralizationand, upon formation, does not adhere to the steep slope of ribs,but detaches at the rib peak and reattaches towards the posterior,just beyond the foot of the posterior scarps of ribs. In thisway, a sinus or open space developed between the internal surfaceof the periostracum and the outer shell surface along each steeprib slope. These spaces could remain filled with extrapallialfluid after the mantle advances beyond that point during shellsecretion. FGPs grow within this microenvironment, out of contactwith the mantle. Other species with divaricate ribs do not developFGPs simply because the periostracum adheres tightly to both ribslopes (which are never so steep as in Solecurtus and Strigilla).FGPs constitute one of the rare cases of remote biomineralizationin which aragonite is produced and direct contact with the mantlenever takes place. (Received 22 November 1999; accepted 20 February 2000)     

Functional morphology of the mantle of Nautilus pompilius (Mollusca, Cephalopoda)

This study presents histological and cytological findings on the structural differentiation of the mantle of Nautilus pompilius in order to characterize the cells that are responsible for shell formation. The lateral and front mantle edges split distally into three folds: an outer, middle, and inner fold. Within the upper part of the mantle the mantle edge is divided into two folds only; the inner fold disappears where the hood is attached to the mantle. At the base of the outer fold of the lateral and front mantle edge an endo-epithelial gland, the mantle edge gland, is localized. The gland cells are distinguished by a distinct rough endoplasmic reticulum and by numerous secretory vesicles. Furthermore, they show a strong accumulation of calcium compounds, indicating that the formation of the shell takes place in this region of the mantle. Numerous synaptic contacts between the gland cells and the axons of the nerve fibers reveal that the secretion in the area of the mantle edge gland is under nervous control. The whole mantle tissue is covered with a columnar epithelium having a microvillar border. The analyses of the outer epithelium show ultrastructural characteristics of a transport active epithelium, indicating that this region of the mantle is involved in the sclerotization of the shell. Ultrastructural findings concerning the epithelium between the outer and middle fold suggest that the periostracum is formed in this area of the mantle, as it is in other conchiferan molluscs.