Constructional morphology of strombid gastropods

The extreme diversity in shell shape of strombid gastropods is interpreted as the result of three independent factors: (1) The terminal growth pattern of the Strombidae allows the circumvention of geometric constraints on shell morphology found in gastropods with continuous or periodic growth patterns. (2) Shell morphology in the Strombidac is adaptive in epifaunal locomotion, burrowing. infaunal or semi-infaunal habits, and passive protection from predators. Specialization for one of these functions often conflicted with the others. thus bringing about a forced 'choice' among mutually exclusive morphological characters. (3) Conservatism in life habits and anatomy of the soft parts has allowed the multiple evolution of extreme shell morphologies, as well as the secondary return to relativcly unspecialized morphologies. □ Constructional morphology, functional morphology. growth. behaviour. evolution, locomotion, burrowing, predation, exoskeleton. shell. Mollusca. Gastropoda. Strombacea. Strombidae.     

Some morphological adaptations in freshwater bivalves

Savazzi, E. & Yao, P. 1992 04 15: Some morphological adaptations in freshwater bivalves. Lethaia , Vol. 25, pp. 195–209. Oslo. ISSN 0024–1164.
Several freshwater bivalves possess peculiar shell morphologies. An extension of the postero-dorsal shell margins above the hinge line evolved convergently in several unionids. This extension supplements the opening momentum of the ligament, but must be broken off periodically in order to allow further shell growth. Arconaia and Cuneopsis have evolved twisted commissure planes, comparable to those found in unrelated marine bivalves. In marine forms, byssus is believed to have played a fundamental role in the evolution of shell torsion. However, the twisted Unionidae do not possess a byssus in the adult stage, thus forcing us to re-evaluate our ideas on the adaptive value and evolution of shell torsion. Solenaia oleivora is apparently incapable of reburrowing and of retracting its foot within the shell. The foot may be functional as an anchor, and is perhaps involved in chemosynthesis by storing sulphur extracted as sulphide from the surrounding sediment. Other adaptations of freshwater bivalves include selective thickening of portions of the shell that enhance its stability, permanent anterior and posterior gapes, and oyster-like morphologies and shell structures. * Functional morphology, constructional morphology, burrowing, shell torsion, Mollusca, Bivalvia, Unionacea, Recent, Quaternary, People's Republic of China .     

How to recognize in situ fossil cephalopods: evidence from experiments with modern Nautilus

Field and flume experiments with modern Nautilus pompilius establish two prerequisites to recognize in situ preservation of fossil cephalopod shells (soft parts were within body chamber in situ at the time of fossilization): occurrence of the upper jaw within the body chamber and the position of jaws within the body chamber. Morphology of shells and jaws in modern and fossil nautiloids is so similar that these prerequisites can be applied for fossil nautiloids and provide implications for ammonoids. The upper jaws of Nautilus start to move at a water velocity of > 0.2 m/s, when the shells are reoriented with the aperture downstream; jaws are therefore unlikely to be secondarily deposited near the shell aperture by bottom currents. The lower jaws, moved at the velocity of > 0.1 m/s, can be deposited around the shell aperture by weak current (0.1–0.2 m/s in velocity), but never enter the inside of body chamber. Neither jaw is likely to be separately and selectively displaced from the inside of the body chamber through scavenging of the soft parts by burrowing infaunal animals. An upper jaw preserved inside the body chamber, together with a lower jaw, is thus a reliable indicator of in situ preservation; a sole lower jaw preserved around the shell aperture is likely to be secondarily deposited. Sedimentary structures inferring rapid burial events and jaw size are useful as additional evidence. Smaller jaws were more likely to be displaced from the body chamber by scavenging by infaunal animals after in situ burial, so that smaller jaws preserved within the body chamber suggest less scavenging. These findings are crucial to interpreting the taphonomic history and palaeo-ecology of fossil cephalopods.     

Mode of life and soft body shape of heteromorph ammonites

Ebel, K. 1992 04 15: Mode of life and soft body shape of heteromorph ammonites. Lethaia , Vol. 25, pp. 179–193. Oslo. ISSN 0024–1164.
Using the idea of a benthic mode of life for ammonites, based on a gastropod-like shell position, it is possible to reconstruct the development of all heteromorph ammonites by regarding single growth stages and the presumable acting forces. The reconstruction of shell formation, particularly the final shell position of heteromorphs with a hook, indicates that the soft body of the ammonite animal was considerably larger than comparison with the present-day Nautilus would suggest.     

Geometric shell growth and mode of life of a Devonian chonetoidean brachiopod

Morphological and statistical analysis of the chonetoid species Kentronetes variabilis from the Lower Devonian (Lochkovian) of the Argentine Precordillera demonstrate ontogenetic changes and allometric relationships between characters. A special study was made of spine distribution, morphology, and growth, compared to valve growth. The first, inner, developed spines (pairs 1–1'and 2–2') continued to grow after development of the following outer pairs. The spacing of spines, their diameter, and the density of growth rings vary from beak to posterolateral margins following a specific 2ⁿ geometric growth factor, compared to the regular, almost linear growth of the valves, attested by growth lines. The linear growth rate of outer spines (pairs 3–3'and 4–4') can be 6–8 times more rapid than that of the shell on the valve margin. Ontogenetic changes in spine morphology are interpreted as a response to changes in the mode of life.     

Opisoma excavatum Boehm,a Lower Jurassic photosymbiotic alatoform‐chambered bivalve

Alatoform bivalves are a polyphyletic group characterized by antero‐posteriorly compressed shells and a ventro‐lateral wing originating from a tight fold of the shell wall. This bizarre shell morphology is interpreted as an adaptation for algal photosymbiosis in heliophilous bivalves. The group contains the living heart cockle Corculum together with four extinct genera ranging in age from the Permian to the Jurassic. The Jurassic alatoform bivalve is Opisoma, which has an aragonitic shell that is divided into two regions, both with different functions: one for stabilization, the other for hosting symbionts. The dorsal part of the shell is massive and played the stabilization role. The ventral part has a very thin and fragile shell that permitted the transmission of light into the internal tissues harbouring photosymbionts. The morphology of this delicate ventral part has thus far remained obscure, due to lack of preservation. Accumulations of Opisoma excavatum Boehm with exquisitely preserved shells containing the fragile winged ventral parts are common within the Pliensbachian shallow‐water, lagoonal carbonate succession of the Rotzo Formation of northern Italy. The wings have internal curved chambers limited by septa parallel to the wing edge. The shell of the ventral part consists of irregular fibrous prismatic and homogeneous structures which progressively infill the chambers. As the chambered wings are analogous structures among alatoform bivalves, they are no longer considered a taxonomic character. According to the observed shell orientation in the field and the consequent organization of the soft parts, Opisoma had an opisthogyrate shell.     

Rare Mollusca from the Lower Devonian Hunsrück Slate of southern Germany

Only ten gastropods have been recovered and a single hyolith found in the radiographic study of the Hunsrück fauna. Four gastropods are interpreted as retaining soft parts: the radula may be identified in one and perhaps the liver and digestive organs in the others. These gastropods may have moved into the area on floating vegetation. The hyolith probably lived on the bottom. Soft parts are present within its shell, but they are not readily interpreted. A tentacular mass anterior to a possible mantle cavity is present, with the bulk of the shell being filled by enigmatic tissue. ▭ Gastropoda, Hyolitha, soft parts, radula, pyritization.     

Segmental growth in planulate ammonites: inferences on costal function

Checa, Antonio & Westermann, Gerd E. G. 1989 01 15: Segmental growth in planulate ammonites: inferences on costal function. Lethaia , Vol. 22, pp. 95–100. Oslo. ISSN 0024–1164.
In planulate Ammonitina, the directions of costae and their parts coincide with the growth lines, both being accreted in parallel and subtriangular increments, i.e. segments. This implies that the growing margin was consistently corrugated (plicated) parallel to the edge of the peristome. The adventral separation of the ribs caused by the coiling increased with whorl involution and was compensated for by costal division into primaries and secondaries. We propose that costation of the shallow-water, nekto-benthic plandates reduced the stiffness and increased effective thickness of their growing peristome. Ribs functioned (primarily or secondarily) as a protection for the immature, fragile aperture against predators and impacts.□ Ammonite shell, Jurassic, morphology, growth lines, ornamentation, costae .     

First record of colour polymorphism in the Neotropical apple snail Asolene platae: inheritance mechanism and evidence for multiple paternity

Asolene platae (Ampullariidae) is a dioecious freshwater snail with subaquatic gelatinous egg masses that dwells in the Río de la Plata basin (Argentina). The aim of this study was to describe the inheritance mechanism of the colour variations of the shell and soft parts of this snail, and to study their potential use as a genetic marker. The wild-type phenotype presents dark pigments in the soft parts and in shell bands, whereas the yellow phenotype lacks dark pigments in the soft parts and also most dark bands in the shell, except for a subsutural and a periumbilical band. The data showed that the lack of pigments in A. platae is a recessive homozygotic condition with a simple Mendelian inheritance mechanism. Females of the wild-type phenotype had a higher number of bands than the males. The pigment of the bands of both phenotypes is located in the calcareous matrix of the shell. Using the lack of pigments as a genetic marker we demonstrated the existence of biparental egg masses in A. platae, hitherto known in only one species within the Ampullariidae.     

Burrowing mechanisms and sculptures in Recent gastropods

Savazzi, Enrico 1989 01 15: Burrowing mechanisms and sculptures in Recent gastropods. Lethaia , Vol. 22, pp. 31–48. Oslo. ISSN 0024–1164.
Burrowing was observed in 32 gastropod species, belonging to 8 families, from Italy and the Philippines. Most species burrow by repeating a three-phase sequence: (1) digging with the foot, (2) dragging the shell forward and downward, and (3) rocking the shell around its longitudinal axis. Minor specific differences in the burrowing dynamics are common, and totally different mechanisms also occur. Burrowing sculptures consisting of terraces or asymmetrical tubercles are observed in the majority of the studied species. Characteristics of the burrowing process explain some cases of apparent divergence of burrowing sculptures from the paradigm. Burrowing sculptures in the Gastropoda should be expected to occur mostly among medium-slender, rather than markedly high-spired, shell morphologies. □ Mollusca, Gastropoda, Cerithiidae, Nassariidae, Mitridae, Costellariidae, Conidae, Terebridae, Turridae, Hydro-biidae, shell, sculpture, burrowing, functional morphology, ecology, behaviour, Holocene, Indo-Pacific. Philippines, Mediterranean, Italy .     

Observations on the shell morphology of two species of cockle Cerastoderma edule and C. glaucum

Aspects of the shell morphology of the cockles Cerastoderma edule (L.) and C. glaucum (Poiret) collected from the R. Crouch and R. Roach (Essex, England) are considered with reference to interpretation of the environment of sub-fossil assemblages. Discrimination between the two cockle species is best achieved by assessment of qualitative shell characters, quantitative measurements being of rather limited value. Factors influencing shell rib-number differ in the two species of Cerastoderma; within a common environment C. glaucum has fewer ribs than C. edule . Mean rib-number of C. edule is directly related to salinity over the colonized salinity range 20–36.5%₀ whilst in C. glaucum rib-number is lowest in both low (<10%₀) and high (>35%₀) salinity environments, being maximal where salinities between 20–30%₀ occur. Internal shell pigmentation may be absent in C. glaucum from low salinity environments. Magnesium content of whole shells is similar in the two cockle species from a mixed population. In both cockles, however, the prismatic layer contained more magnesium than the nacreous shell layer. The most useful molluscan indicators of brackish water lagoon environments are C. glaucum and Hydrobia ventrosa; these species being absent from areas of only limited wave action. C. edule may be associated with marine or estuarine molluscan indicators.     

Sectorial expansion and shell morphogenesis in molluscs

Checa, A. 1991 01 15: Sectorial expansion and shell morphogenesis in molluses. Lethaia . Vol. 24, pp. 97–114. Oslo. ISSN 0024–1164.
Any coiled shell can be described as a series of independent helicospirals that join homologous points along the shell surface. The cross-section is therefore seen as a set of points. obtained at its intersection with the helicospirals. Any cross-sectional sector contained between two adjacent points is capable of expanding or contracting during the development and a differential parameter ( L'/L ) has been devised to quantify this expansion rate. The morphometrics so obtained is here called sectorial expansion analysis. This analysis and other related procedures have been applied on cross-sections and apertures in selected Molluxa with the aim of evaluating the incidence of sectorial expansions on shell shape. Those parameters affecting whorl expansion rate and whorl overlapping may be directly modified by sectorial expansions. Changes in the mode of coiling (curvature and torsion) often, but not always, involve sectorial expansions, perhaps as a constructional feature. This approach reveals the advantages and drawbacks of the present analysis as compared to previous theoretical models. Sectorial expansion. morphogenesis. ornamentation, septal suture, coiled shell, gastropods, bivalues. ammonites .     

The shell-structure, growth and functional morphology of some Lower Devonian athyrids from northwest Spain

Alvarez, F. & Brunton, C. H. C. 1990 04 15: The shell-structure, growth and functional morphology of some Lower Devonian athyrids from northwest Spain. Lethaia , Vol. 23, pp. 117–131. Oslo. ISSN 0024–1164.
In the Emsian (Lower Devonian) of northwest Spain three congeneric species of athyrids have been recognized principally by their, as yet, unique shell microstructures. The shell remains thin throughout ontogeny, with hardly any internal thickening, while the external lamellac are long and two to four times as thick as the valves. being composed of many layers of secondary fibres. A growth model for these shells is presented, including the recognition and differentiation of true valve growth as compared to growth of lamellae. Functional morphology is discussed and a new explanation presented for the function of the visceral foramen. The possible growth and disposition of the lophophore are described and, taken together with morphological and environmental factors, an anterior inhalant water circulation system is proposed. * Brachiopods, Athyrididae, Devonian, shell-structure, growth. functional morphology .     

Effects of foliaceous varices on the mechanical properties of Chicoreus dilectus (Gastropoda: Muricidae)

Removal of the foliaceous varices from the shells of Chicoreus dilectus had no significant effect on the crushing strength (load at failure) of the shells, but decreased the total work necessary to break the shells by 35–50%. This effect was due to the removal of varices as an architectural feature of the shell rather than simply to the removal of shell material per se ; varicate shells had a mean work to failure per unit shell mass 1–6–2–3 times greater than avaricate shells. Because the strength of gastropod shells often exceeds the maximum forces that durophagous predators may bring to bear on them, work to failure may be a more important factor in the design of gastropod shells than breaking load.     

A new lacazelline species (Brachiopoda,Recent) from the Maldive Islands,Indian Ocean

A new lacazelline brachiopod species is described from reef caves in the outer reef slope zone off Addu and South Male atolls in the Maldive Islands, Indian Ocean. Based on anatomical features of the soft parts, shell morphology and shell microstructure the new taxon is assigned to the genus Ospreyella Lüter and Wörheide 2003 in the subfamily Lacazellinae of the family Thecideidae. This new species, here named O. maldiviana and representing the first occurrence of the genus in the Indian Ocean, is compared with the type species O. depressa Luter from Osprey Reef, Coral Sea, Australia. Comparisons are made between Ospreyella (now two species) and the other extant lacazelline genera Lacazella Munier‐Chalmas (three species) and Pajaudina Logan (one species).     

How did the carrier shell Xenophora crispa (König, 1825) build its shell? Evidence from the Recent and fossil record

The genus Xenophora comprises species of marine gastropods (Cretaceous-Recent) able to add fragments of various origins to their shell surface. Agglutination potentials vary, from species lacking attachments to species completely covered by agglutinated materials, as in the Mediterranean species Xenophora crispa. Here, we analyse Recent and fossil specimens of Xenophora crispa from the Mediterranean area using SEM and XRD, to better understand their biomineralization patterns and the mechanisms leading to the agglutination of shells, bioclasts and lithoclasts, and their evolution in time. We also provide new data on poorly studied gastropod shell microstructures. We conclude that: (1) most of the Xenophora crispa shell consists of an aragonitic crossed lamellar fabric, but fibrous to spherulitic prismatic fabrics, seemingly of calcite, have been found in the columella and peripheral edge (the thickest parts of the shell); (2) attachment of objects is mediated by a prismatic microstructure, indicating that this may be the most functional fabric in attachment areas in molluscs; and (3) the functional purpose of the agglutination in Xenophora crispa may be related to a snowshoe strategy to successfully colonize muddy substrates, coupled with tactile and olfactory camouflage. Indeed, this species secretes in the columella and peripheral edge a less dense and a more organic rich calcitic fabric, possibly to lighten the shell thickest parts in order not to sink in soft sediments and to facilitate the shell raising from the substrate to create a protected feeding area. This behaviour seems to have been maintained by X. crispa over 2 My time span.     

Prolonged morphometric study of barnacles grown on soft substrata of hydrogels and elastomers

A long-term investigation of the shell shape and the basal morphology of barnacles grown on tough, double-network (DN) hydrogels and polydimethylsiloxane (PDMS) elastomer was conducted in a laboratory environment. The elastic modulus of these soft substrata varied between 0.01 and 0.47 MPa. Polystyrene (PS) (elastic modulus, 3 GPa) was used as a hard substratum control. It was found that the shell shape and the basal plate morphology of barnacles were different on the rigid PS substratum compared to the soft substrata of PDMS and DN hydrogels. Barnacles on the PS substratum had a truncated cone shape with a flat basal plate while on soft PDMS and DN gels, barnacles had a pseudo-cylindrical shape and their basal plates showed curvature. In addition, a large adhesive layer was observed under barnacles on PDMS, but not on DN gels. The effect of substratum stiffness is discussed in terms of barnacle muscle contraction, whereby the relative stiffness of the substratum compared to that of the muscle is considered as the key parameter.     

Shell cracking strength in almond (Prunus dulcis [Mill.] D.A. Webb.) and its implication in uses as a value-added product

Researchers are currently developing new value-added uses for almond shells, an abundant agricultural by-product. Almond varieties are distinguished by processors as being either hard or soft shelled, but these two broad classes of almond also exhibit varietal diversity in shell morphology and physical characters. By defining more precisely the physical and chemical characteristics of almond shells from different varieties, researchers will better understand which specific shell types are best suited for specific industrial processes. Eight diverse almond accessions were evaluated in two consecutive harvest seasons for nut and kernel weight, kernel percentage and shell cracking strength. Shell bulk density was evaluated in a separate year. Harvest year by almond accession interactions were highly significant (p  0.01) for each of the analyzed variables. Significant (p  0.01) correlations were noted for average nut weight with kernel weight, kernel percentage and shell cracking strength. A significant (p  0.01) negative correlation for shell cracking strength with kernel percentage was noted. In some cases shell cracking strength was independent of the kernel percentage which suggests that either variety compositional differences or shell morphology affect the shell cracking strength. The varietal characterization of almond shell materials will assist in determining the best value-added uses for this abundant agricultural by-product.     

Higher risk of fatality by predatory attacks in earlier ontogenetic stages of modern Nautilus pompilius in the Philippines: evidence from the ontogenetic analyses of shell repairs

The shell repair scars of modern Nautilus pompilius in the Philippines presumably due to sub‐lethal predatory attacks were examined throughout ontogeny ranging from hatching to maturity, revealing the higher risk of fatality in earlier ontogenetic stages. The examinations throughout ontogeny demonstrate that: (1) sub‐lethal predatory attacks by other organisms have no preferred position through ontogeny; (2) the sizes of shell repair scars are similar throughout ontogeny and, therefore, the width and length of shell repair scars relative to shell diameter decrease from hatching towards maturity; and (3) the proportion of sub‐lethal predatory attacks within 10 mm increments of shell diameter are similar irrespective of shell diameter or decrease approaching maturity. The numbers of the irregular, radiating, black shell repair scars, indicating injury to the soft parts at shell breakage, decrease at a diameter >80 mm and none were found at <56 mm in shell diameter. Furthermore, the rarity of juvenile trapping data might be related to the higher fatality from predatory attacks in earlier ontogenetic stages, providing crucial information for the conservation of N. pompilius.