The effect of age at metamorphosis on the transition from larval to adult suspension‐feeding of the slipper limpet Crepidula fornicata

Slipper limpets use different ciliary feeding mechanisms as larvae and adults. Veliger larvae of Crepidula fornicata developed part of the adult feeding apparatus, including ctenidial filaments, neck lobe, and radula, before metamorphosis, but ctenidial feeding did not begin until well after loss of the larval feeding apparatus (velum) at metamorphosis. Earlier initiation of ctenidial feeding by individuals that were older larvae when metamorphosis occurred suggests continued development toward ctenidial feeding during delay of metamorphosis. Early juveniles produced a ciliary current through the mantle cavity and moved the radula in a grasping action before they began to capture algal cells on mucous strands or form a food cord. Either early juveniles could not yet form mucous strands or they delayed their production until development of other necessary structures. The neck canal for transporting food from ctenidium to mouth cannot develop before velar loss. In their first feeding, juveniles fed much like the adults except that the neck canal was less developed and the path of the food cord toward the mouth sometimes varied. As suspension feeders, calyptraeids lack the elaborations of foregut that complicate transition to juvenile feeding for many caenogastropods, but a path for the food cord must develop after velar loss. Why individuals can initiate ctenidial feeding sooner when they are older at metamorphosis is not yet known. The juveniles became sedentary soon after metamorphosis and were not observed to feed by scraping the substratum with the radula, in contrast to the first feeding by juveniles of another calyptraeid species, observed by Montiel et al. ( 2005 ).     

The biology of Dendropoma corallinaceum and Serpulorbis natalensis, two South African vermetid gastropods

D. corallinacewn forms sheet-like colonies at Mean Low Water (MLW) on exposed rocky shores: S. natalensis forms loose aggregations under stones below MLW in more sheltered situations. Both species released young from July— December. D. corallinaceun has single embryos in capsules brooded free in the mantle cavity; S. natalensis has 20–40 embryos per capsule, the latter being attached to the roof of the shell and hanging through a dorsal slit in the mantle. The morphology of the crawling young is similar in both species, resembling adult errant prosobranchs. D. corallinaceum has a stouter, more streamlined protoconch than S. natalensis correlated with the more turbulent water of its habitat. The crawling young secrete a sticky mucous trail for adhesion to the substratum. They are strongly photopositive until settlement. D. corallinaceum can settle within 24 hours but may delay settlement to 4–5 days if suitable substrata are absent. Metamorphosis is complete 2 days after settlement. D. corallinaceum prefer to settle on Lithothamnion which is easily eroded by the radula to provide a groove for rapid, firm attachments to the substratum. Both species feed from mucous nets, S. natalensis having a slower feeding rhythm commensurate with its larger size. Contiguous pairs of S. natalensis have synchronised feeding rhythms, probably to reduce mutual net robbing. Particles are also filtered out of suspension; many particles are rejected in the pseudofaeces but some are ingested. D. corallinaceum and S. natalensis have many similar ecologically equivalent species throughout the warm temperate and tropical seas of the world.     

The influence of season and the tidal cycle on division of labour by the radula during feeding in the estuarine brooding gastropod <Emphasis Type="Italic">Crepipatella dilatata</Emphasis> (Calyptraeidae)

The brooding gastropod Crepipatella dilatata can feed by scraping the substrate with the radula and by suspension-feeding, which also requires use of the radula. There is a “division of labour” for the radula among three discrete tasks associated with feeding: (1) removing mucous balls from the food pouch; (2) transferring the mucous cord from the neck channel to the mouth (both components of suspension-feeding); (3) scraping the substrate. We hypothesised that the proportion of time used for each feeding activity varies according to environmental conditions. Total radular activity in females was greatest at high tide and in summer. The rate of radular extrusion for ingesting the mucous cord varied seasonally and between brooding and non-brooding females. Non-brooding females exhibited higher rates of radular extrusion for ingesting the mucous cord and for scraping the substrate than did brooders. In females, radular activity in removing the mucous ball from the food pouch was strongly influenced by the tidal cycle during winter, reaching minimum values at low tide. Differences were recorded in substrate scraping among seasons and within tidal cycles, and among males, brooding females and non-brooding females. Brooding females displayed less rasping than non-brooders, since the area available for grazing was restricted by the egg mass. Throughout the year, including low salinity periods, males allocated a greater proportion of total radular activity to rasping than to removing the mucous ball or ingesting the mucous cord. The feeding behaviour of both males and females is modulated by salinity, but the principal determinants of radular activity are the mode of reproduction (brooding in females) and, in males, motility.     

Further studies on the morphology of the Triviidae (Gastropoda: Prosobranchia) with emphasis on species from southern Africa

Aspects of the morphology of II species of Triviidae are described, including the living animal, shell, radula, mantle complex, central nervous system and reproductive system. Interspecific differences in radular morphology are important in distinguishing closely allied species. Trivia aperta, T. costata and T. verhoeft are protandric hermaphrodites while T. caluariola, T. neglccta, T. pellucidula, T. rubra, T. solandri and T. suavis are gonochoric. The elaboration of the receptaculum seminis varies between species. In T. aperta, T. calvariola, T. neglecta and T. who& the receptaculum is undivided while in T. costafa, T. pellucidula, T. solandri and T. suavis there are two or more lobes of the receptaculum.     

Deposit feeding in Abra tenuis (Bivalvia: Tellinacea)

Deposit feeding in Abra tenuis is described in terms of the size of particles utilized.
Material is collected by the inhalant siphon performing a circular motion sucking in sediment from beneath and on the surface.
The size distribution of silica admitted into the mantle cavity is described and shown to be controlled by physical parameters. The density of a particle does not affect its uptake by the inhalant siphon. The size distribution of the sediment affects the size distribution of particles admitted to the mantle cavity.
No selection of material for ingestion in terms of size occurs after it has been taken into the mantle cavity. Thus the range of material ingested is ultimately controlled by a physical parameter, the inhalant opening.     

Some aspects of feeding in thecosomatous pteropod molluscs

Various aspects of the feeding behavior of thecosomatous pteropods of the families Cavoliniidae and Cymbuliidae have been examined both in the field with SCUBA, and in the laboratory. It was found that the family Cavoliniidae employs a ciliary-mucus feeding mechanism in which the mantle cavity water current and pallial gland are of primary importance in food collection. The feeding mechanism is discussed, as well as related aspects including buoyancy control, particle size range of natural diets, oxygen consumption, and water filtration rates. It is concluded that Cavolinia must have a high efficiency of retention and assimilation of available carbon.     

Studies on the in vitro formation of periostracum inHelisoma duryi: the influence of brain

Summary Excised mantle tissue produces periostracum when placed in organ culture. Mantle collars taken from animals exhibiting a fast rate of shell deposition in vivo produce more periostracum than mantle collars from slow growing animals. The addition of a brain from a fast growing animal increased periostracum production by a mantle collar from a slow growing animal. This effect was enhanced by removing the cerebral ganglia lateral lobes. This suggests that a factor(s) is present in the brain of fast growing animals which enhances periostracum formation in vitro. The lateral lobes appear to inhibit this activity. Radiolabel incorporation studies suggest that the periostracum produced by fast growing animals has a higher glycine content than that produced by slow growing animals.Abbreviations Dopa 3,4-dihydroxyphenylalanine - FB brain derived from a fast growing animal - FMC mantle collar derived from a fast growing animal - LL lateral lobes - LL lateral lobes removed - MC mantle collar - SB brain derived from a slow growing animal - SGPF shell growth promoting factor - SMC mantle collar derived from a slow growing animal - TCA trichloroacetic acid - WM whole mantle     

Feeding adaptations of Melagraphia aethiops (Gmelin), an intertidal trochid mollusc

The feeding behaviour of the intertidal trochid Melagraphia aethiops (Gmelin) on a sheltered stony shore was investigated. The effects of snail size, season, and shore level on feeding were examined. Feeding was examined in the laboratory in aquaria fitted with glass plates coated with algal and detrital films. Rates of radula rasping and the frequency of feeding bouts (feeding activity) were measured.

Radula rate varied little with snail size and large snails were 35% more active feeders than small snails. Small snails, however, consumed three to four times as much food per radula rasp, on a unit weight basis, as large snails. Winter acclimatization involved an increased tolerance of low water temperatures and an overall 60% decrease in radula rasping rate compared with summer acclimatized animals. Concurrently, field crawling rates decreased 71% during the summer-winter transition suggesting that lower food intake was balanced by reduced locomotory activity. High and low shore inhabiting snails showed little difference in radula rasping rates. High shore snails, however, compensated for their shorter feeding periods by being 37% more active in feeding and crawling in the field 40% faster than low shore snails.     

The anatomy of the opisthobranch genus Hydatina and the functioning of the mantle cavity and alimentary canal

A study of specimens of Hydatina physis (Linne, 1758) from Australia, New Zealand and Hawaii shows that the radula is variable and that sometimes a central tooth is present. A comparison with a specimen from the Canary Islands suggests thatH. stromfelti Odhner, 1932 is a geographical form of H. physis. A similar explanation applies to the small shell variations found in West Indian forms, sometimes described as H. vesicaria Solander, 1786.
The anatomy of specimens of Hydatina amplustre (Linne, 1758) confirms that this species is congeneric with H. physis. It is suggested that members of this genus feed specifically on cirratulid polychaetes. The nervous system, reproductive system and mantle cavity show that the Hydatinidae are primitive opisthobranchs, closely related to the Acteonidae and the Bullinidae.     

Neuroendocrine Control Mechanisms in Shell Formation

The brain of Helisoma duryi contains several neurodendocrinecentres. Factors) present in the cerebral ganglia are thoughtto be involved in normal shell growth while neurosecretory substancespresent in the visceral ganglion are involved in the repairof damaged shell. In Lymnaea stagnalis a growth hormone is producedby the cerebral ganglion which stimulates periostracum formationand the calcification of the inner shell layer. The second effectis thought to occur through the action of a mantle edge calciumbinding protein. In Helisoma, mantle collar is able to produce the periostracumin vitro. The presence of brain from a fast growing donor increasesthe amount of periostracum produced by a mantle collar froma slow growing animal. This effect is further enhanced by theremoval of the lateral lobes. The periostracum produced by fastgrowing animals has a higher glycine content than that producedby slow growing snails. The presence of dorsal epithelial tissueenhances the incorporation of calcium into periostracum formedin vitro. These findings suggest that a single factor is present in thebrain of fast growing Helisoma which modulates shell formationrates in vivo and periostracum formation in vitro.     

Main patterns of radula formation and ontogeny in Gastropoda

Gastropoda is morphologically highly variable and broadly distributed group of mollusks. Due to the high morphological and functional diversity of the feeding apparatus gastropods follow a broad range of feeding strategies: from detritivory to highly specialized predation. The feeding apparatus includes the buccal armaments: jaw(s) and radula. The radula comprises a chitinous ribbon with teeth arranged in transverse and longitudinal rows. A unique characteristic of the radula is its continuous renewal during the entire life of a mollusk. The teeth and the membrane are continuously synthesized in the blind end of the radular sac and are shifted forward to the working zone, while the teeth harden and are mineralized on the way. Despite the similarity of the general mechanism of the radula formation in gastropods, some phylogenetically determined features can be identified in different phylogenetic lineages. These mainly concern shape, size, and number of the odontoblasts forming a single tooth. The radular morphology depends on the shape of the formation zone and the morphology of the subradular epithelium. The radula first appears at the pre- and posttorsional veliger stages as an invagination of the buccal epithelium of the larval anterior gut. The larval radular sac is lined with uniform undifferentiated cells. Each major phylogenetic lineage is characterized by a specific larval radula type. Thus, the docoglossan radula of Patellogastropoda is characterized by initially three and then five teeth in a transverse row. The larval rhipidoglossan radula has seven teeth in a row with differentiation into central, lateral, and marginal teeth and later is transformed into the adult radula morphology by the addition of lateral and especially marginal teeth. The taenioglossan radula of Caenogastropoda is nearly immediately formed in adult configuration with seven teeth in a row.     

Mantle histology and histochemistry of three pearl oysters: Pinctada persica,Pinctada radiata and Pteria penguin

Shells in pearl oysters are produced by the mantle which is also used as a graft in pearl operations. Here, we investigate the mantle structure of a new pearl oyster species of the Persian Gulf, Pinctada persica, and compare it to two other pearl-producing species, Pinctada radiata and Pteria penguin. The anterior, ventral and posterior segments of the mantle edge of each valve were fixed, and tissue sections were stained with haematoxylin and eosin. A new pentachrome method and PAS-alcian blue staining were used to characterise the different mucosubstances. The mantle edges were found to have an outer, middle and inner fold, which have different morphology in each species. The mantle edge is lined by cuboidal and columnar epithelia, and interspersed among these epithelial cells we found mucous cells and cells that contained brown granules. The outer and middle folds of the two Pinctada species show different shapes to that of Pteria penguin. Most of the mucous cells in the mantle contain acidic mucosubstances and small amounts of mixed acidic-neutral mucosubstances were observed in the middle and inner fold of Pinctada persica. This study reveals that the mantle edges of the three species possess similar cellular structure, but vary in the shape of the folds.     

Burrowing of the lingulid brachiopod Glottidia pyramidata: its ecologic and paleoecologic significance

Contrary to popular assumption, the pedicle of Glottidia is not its principal burrowing organ. The brachiopod props itself up with the pedicle and enters the sediment with the valves leading, anterior end first. The pedicle trails behind. Burrowing is accomplished by cyclical valve motions: rotary, sliding, and gaping movements are used. Rapid valve closure ejects water from the mantle cavity to loosen the sediment. The lateral setae convey mucus-bound sand posteriorly (upwards). X-radiography shows that the burrows are U-shaped: in a few hours, the animal reappears in feeding position with the anterior end pointing out of the sediment and the pedicle extending down into the burrow. This burrowing process explains the substrate preferences of lingulids. The thick. closely spaced setae are adapted for burrowing. The spacing between individuals is increased, suggesting competition for food.     

A New Genus and Species of the Family Anulidentaliidae (Scaphopoda: Dentaliida) and its Systematic Implications

Shell, radula, and anatomy of Epirhabdoides ivanovi new genusand species are described from a sample of the Russian VitjazExpedition from the Japan Trench. It is distinguished from thesimilar Laevidentalium sominium by shell morphometrics and radulamorphology. The radula is almost identical with that of Anulidentaliumbambusa (Anulidentaliidae). The anatomy of the mantle margins,however, including dorsolateral slits at the anterior mantlemargin and a connective tissue bolster at the posterior mantleopening is that of the family Rhabdidae. In a parsimony analysisEpirhabdoides ivanovi takes an intermediate position betweena basal grade of Gadilinidae and the remaining Dentaliida implyingconvergent evolution of mantle characters. An alternative butless parsimonious tree with E. ivanovi as sister taxon to Rhabdusrequires convergences in radula characters. This is the firstdocumented case of convergent anatomical features among Scaphopodaand enhances the need of radula and soft part investigationof the conchologically little informative, smooth-shelled dentaliidgroups. (Received 16 March 1998; accepted 1 June 1998)     

Self-assemblage formation in a social insect: the protective curtain of a honey bee swarm

Summary. This paper considers a little-studied topic in the biology of social insects: the formation of self-assemblages. It focuses on the mechanisms whereby the outermost workers in a bivouacked swarm of honey bees, when rained upon, form a water repellent curtain of bees over the swarm cluster. Specifically, we analyzed how the worker bees in the mantle of a swarm cluster adjust their body orientation, wing spread, and inter-individual spacing to form a protective curtain when wetted. When warm and dry, the mantle bees orient their bodies weakly with respect to gravity, do not tuck their heads under adjacent bees, have high variability in wing spread, and space themselves widely. In contrast, when warm and wet, the mantle bees orient uniformly with head upward, tuck their heads beneath the abdomens of bees above, hold their wings together, and press tightly together. This produces a surface that closely resembles a tiled roof. When cool and dry, the mantle bees generally orient their bodies with head upward, press their heads into the interior of the cluster, hold their wings wide apart, and draw close together. We also examined the age distribution of the mantle bees. Older bees are more likely than younger bees to be found in the mantle of a swarm, perhaps because younger bees are more important than older bees to colony survival after swarming and so occupy a more sheltered position in a swarm. Finally, we tested whether swarm clusters that have formed a protective curtain shed water more effectively than ones that have not formed a curtain. We found that this is the case.Received 28 November 2003; revised 29 February 2004; accepted 11 April 2004.     

Functional morphology of the mammiliform penial glands ofLittorina saxatilis (Gastropoda)

Summary The functional morphology of the mammiliform penial glands ofLittorina saxatilis has been investigated with both light and electron microscopy. These penial glands line the ventral edge of the penis and orient with the female mantle during copulation. Secretions are released from the penial glands to this interface where they probably function in adhesion. The penial gland secretions comprise heterogeneous granules as well as apocrine and mucous secretions. The heterogeneous granules are produced in separate multicellular glands arranged in a series of lobes that lie outside a thick smooth muscle layer enclosing the lumen. Each glandular lobe is surrounded by a thin layer of smooth muscle. Secretions are transported in individual cellular processes that pass through the thick smooth muscle layer and empty into the lumen. Surrounding the lumen is an epithelium containing apocrine secretory cells as well as occasional goblet-type, mucous cells. The combined action of the muscles forces secretions out of the lumen through the penial papilla, onto the external surface of the mammiliform penial gland. Longitudinal muscles extend into the penial papilla enabling its protrusion or retraction. Retraction of the penial papilla following secretion release is thought to create negative pressure beneath the penial gland producing suction adhesion. The visco-elastic properties of the penial gland secretion are qualitatively different from foot mucus and may represent specialization to an adhesive function.     

Original molluscan radula: comparisons among Aplacophora,Polyplacophora, Gastropoda,and the Cambrian fossil Wiwaxia corrugata

As the original molluscan radula is not known from direct observation, we consider what the form of the original radula may have been from evidence provided by neomenioid Aplacophora (Solenogastres), Gastropoda, Polyplacophora, and the Cambrian fossil Wiwaxia corrugata (Matthews). Conclusions are based on direct observation of radula morphology and its accessory structures (salivary gland ducts, radular sac, anteroventral radular pocket) in 25 species and 16 genera of Aplacophora; radula morphogenesis in Aplacophora; earliest tooth formation in Gastropoda (14 species among Prosobranchia, Opisthobranchia, and Pulmonata); earliest tooth formation in four species of Polyplacophora; and the morphology of the feeding apparatus in W. corrugata. The existence of a true radula membrane and of membranoblasts and odontoblasts in neomenioids indicates that morphogenesis of the aplacophoran radula is homologous to that in other radulate Mollusca. We conclude from p redness of salivary gland ducts, a divided radular sac, and a pair of anteroventral pockets that the plesiomorphic state in neomenioids is bipartite, formed of denticulate bars that are distichous (two teeth per row) on a partially divided or fused radula membrane with the largest denticles lateral, as occurs in the genus Helicoradomenia. The tooth morphology in Helicoradomenia is similar to the feeding apparatus in W. corrugata. We show that distichy also occurs during early development in several species of gastropods and polyplacophorans. Through the rejection of the null hypothesis that the earliest radula was unipartite and had no radula membrane, we conclude that the original molluscan radula was similar to the radula found in Helicoradomena species.     

Regional specification during embryogenesis in the inarticulate brachiopod Discinisca.

The process of embryogenesis is described for the inarticulate brachiopod Discinisca strigata of the family Discinidae. A fate map has been constructed for the early embryo. The animal half of the egg forms the dorsal ectoderm of the apical and mantle lobes. The vegetal half forms mesoderm and endoderm and is the site of gastrulation; it also forms the ectoderm of the ventral regions of the apical and mantle lobes of the larva. The plane of the first cleavage goes through the animal-vegetal axis of the egg along the future plane of bilateral symmetry of the larva. The timing of regional specification in these embryos was examined by isolating animal, vegetal, or lateral regions at different times from the 2-cell stage through gastrulation. Animal halves isolated at the 8-cell and blastula stages formed an epithelial vesicle and did not gastrulate. When these halves were isolated from blastulae they formed the cell types typical of apical and mantle lobes. Vegetal halves isolated at all stages gastrulated and formed a more or less normal larva; the only defect these larvae had was the lack of an apical tuft, which normally forms from cells at the animal pole of the embryo. When lateral isolates were created at all developmental stages, these halves gastrulated. Cuts which separated presumptive anterior and posterior regions generated isolates at the 4-cell and blastula stages that formed essentially normal larvae; however, at the midgastrula stage these halves formed primarily anterior or posterior structures indicating that regional specification had taken place along the anterior-posterior axis. The plane of the first cleavage, which predicts the plane of bilateral symmetry, can be shifted by either changing the cleavage pattern that generates the bilateral 16-cell blastomere configuration or by isolating embryo halves prior to, or during, the 16-cell stage. These results indicate that while the plane of the first cleavage predicts the axis of bilateral symmetry, the axis is not established until the fourth cleavage. The development of Discinisca is compared to development in the inarticulate brachiopod Glottidia of the family Lingulidae and to Phoronis in the phylum Phoronida.     

A possible home for a bizarre Carboniferous animal: is Typhloesus a pelagic gastropod?

By contrast to many previously enigmatic Palaeozoic fossils, the Carboniferous metazoan Typhloesus has defied phylogenetic placement. Here, we document new features, including possible phosphatized muscle tissues and a hitherto unrecognized feeding apparatus with two sets of ca 20 spinose teeth whose closest similarities appear to lie with the molluscan radula. The ribbon-like structure, located well behind the mouth area and deep into the anterior part of the body, is interpreted as being in an inverted proboscis configuration. Gut contents, mostly conodonts, in the midgut area demonstrate that Typhloesus was an active predator. This animal was capable of propelling itself in the water column using its flexible body and a prominent posterior fin. The affinity of Typhloesus as a pelagic mollusc remains problematic but may lie more closely with the gastropods. Heteropod gastropods share with Typhloesus an active predatory lifestyle and have a comparable general body organization, albeit they possess characteristic aragonitic shells and their origins in the Jurassic post-date Typhloesus. Typhloesus may represent an independent radiation of Mid-Palaeozoic pelagic gastropods.     

Seasonal aberrant radular formation in Thais bronni (Dunker) and T. clavigera (Küster) (Gastropoda : Muricidae)

Radulae of Thais bronni (Dunker) and T. clavigera (Küster) were examined at Mukaishima Island for a period of 2 yr, 1982 to 1984. Radulae of both species are similar in morphology, both having the basic pentacuspid rachidian plan. Sexual dimorphism of the radula was not observed, but rachidian tooth changes morphologically in different growth stages. Seasonal conditions affect the size and shape of the radula; in winter it is clearly malformed and strikingly thin. These aberrant parts of the radula comprised some dozens of rows, in which only several extremely thin rows exist. Results of experiments using T. clavigera under different water temperature conditions showed that the radula is rarely produced below 10 °C and that rate of radular production and replacement increases with increase in temperature. These results suggest that in the field the radula of these species is replaced entirely 2–2.5 times per year and 10–15 times during the life of the animal.