Anatomy and systematic affinity of Stanleya neritinoides (Smith, 1880), an enigmatic member of the thalassoid gastropod species flock in Lake Tanganyika, East Africa (Cerithioidea, Paludomidae)

The radiation of gastropods in Lake Tanganyika is an ideal system for testing competing hypotheses of species flock formation. Yet, much of the basic biology of these species remains unknown. In an ongoing effort to understand the evolution of Tanganyikan gastropods, we here describe Stanleya neritinoides. Alcohol‐preserved material of the soft parts is rare, consequently, the systematic position of the species, and a repeated suggested affinity to Tanganyicia rufofilosa, have been based primarily on features of the shell. However, features of the radula and operculum are unique and do not suggest an affinity to any other Tanganyikan species. Thus, S. neritinoides has remained a particularly poorly known and enigmatic member of the species flock. This investigation confirmed that several aspects of internal and external anatomy are shared between S. neritinoides, T. rufofilosa, and other Tanganyikan gastropods, but that S. neritinoides is unique in features of the radula and seminal receptacle. Moreover, S. neritinoides differs from T. rufofilosa in features of the foregut, midgut, hindgut, kidney, nervous system, reproductive system and reproductive strategy. These new data are inconsistent with an interpretation of identity of Stanleya and Tanganyicia. In addition, given the pervasive differences between the two, a sister‐group relationship between the two is unlikely. More precise systematic placement of S. neritinoides awaits the establishment of a phylogenetic framework for all Tanganyikan gastropods.     

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.     

Functional morphology of vermetid feeding-tubes

Vermetidae are a small family of warm-water sessile gastropods capable of building upright tubes (feeding-tubes) to take advantage of the water flow. Laboratory and field experiments carried out on some Mediterranean species ( Vermetus triquetrus, Vermetus granulatus and Serpulorbis arenaria ) suggest that these structures function as exploratory tubes built not only to receive a better water flow, but mainly to avoid obstacles. In fact, vermetids experimentally exposed in situ to different hydrodynamic conditions do not produce them, but do so in the presence of an obstacle, such as thalli during the massive spring algal growth. This strategy allows them to compete for a virtual space, not directly occupied by the vermetid itself but necessary to spread its mucous net. This interpretation may improve the inference of paleo-environmental events from shell morphology. When building a feeding-tube, vermetids first cut off a portion of the shell on the side towards which they prepare to turn, using their radula, and then produce a new tube formed by short segments, at different angles, till they have reached the desired direction. This process is confirmed by the presence of scars on the shell, composed by a succession of lamellae. The regular distribution of these scars on fossil Petaloconchus intortus , which lived in soft substrates, may be interpreted as a response to periodical anoxic crises or an increase in the sedimentation rates. Their great morphological plasticity makes vermetids close to colonial or modular animals. Thanks to their capability of expressing more than one growth-form, and of re-moulding their shell, they successfully compete for substrate space and are key-stone species in fringe habitats.     

FACTORS AFFECTING PREY CHOICE AND FEEDING TECHNIQUE IN THE CARNIVOROUS SNAIL EUGLANDINA ROSEA FERUSSAC

Euglandina feeds on other gastropods either by quickly suckingthe contents from the shell, or by swallowing the prey whole,which can be more time consuming. When Euglandina were offeredprey species of various sizes, they preferentially consumedthe smaller individuals, and often swallowed these whole. Handlingtime increased with decreasing density since swallowing preywhole became more frequent at the lower densities. The relationship between prey size, predator size, handlingtime and feeding technique was examined. For the two feedingtechniques observed, handling time increased exponentially withthe size of the prey and decreased exponentially with the sizeof the predator. Predators of intermediate size tended to swallowprey whole more frequently than did the smallest and largest. These data are interpreted in terms of the animal behaving insuch a way as to balance its rate of intake of both organicmatter, which resides mainly in the soft parts, and of calcium,which resides mainly in the shell. (Received 11 July 1988; accepted 31 October 1988)     

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.     

A reinterpretation of the mode of life of some Paleozoic frilled gastropods

Linsley, Robert M., Yochelson, Ellis, L. & Rohr, David M. 1978 04 15: A reinterpretation of the mode of life of some Paleozoic frilled gastropods
Most modem gastropods crawl, and support their shells up off the substrate. However. five different groupings of snails predominantly rest their shell upon the substrate. One group, Xenophora- like in habit, lives with the base of the shell propped above the substrate by a frill or spines. Re-examination of the Silurian Euomphalopterus shows that the broad frill is not a selenizone as it has been most frequently interpreted. The broad frill may have served the function of propping the aperture above the substrate. All members of the Euomphalopteridae are reassigned. A review of Paleozoic gastropods suggests that this propping mode of life was adopted more than once by unrelated gastropods.     

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.     

Radula und Fangarme beiMichelinoceras sp. aus dem Silur von Bolivien

New finds of michelinoceratid cephalopods from the Kirusillas-Shale (Ludlow/Silurian) of Ushpa-Ushpa in the Eastern Cordillera of the Bolivian Andes reveal parts of the radula and imprints of arms owing to extremely good preservation conditions. The imprints of soft parts point that the orthoce-rates had 10 arms, two of them shaped to long tentacles as recent coleoids have. Of more importance is the first proof of a michelinoceratid radula in situ. This is the first known ra-dula of lower palaeozoic cephalopods at all. TheMichelinoceras radula consisted of 7 teeth per row. So the radulae of michelinoceratid cephalopods are very similar to those of ammonites and coleoids. But there are great differences in the radula ofMichelinoceras sp. and that ofNautilus sp. All known radulae of fossil and recent cephalopods are compared and phylogenetic or systematic implications are discussed. The classification of the cephalopods into six subclasses as used in the “Treatise” turns out to be unnatural and should be given up. Finally ecological relations between the morphology of radula-elements and mode of life are dis-cussed. Size and number of radula teeth let suppose that thisMichelinoceras specimen must have been an adult individual living far off the coast in pelagic seas.     

Bulinus guernei (Mollusca: Gastropoda) of West Africa: taxonomic status and role as host for schistosomes

The basommatophoran freshwater snail Bulinus guernei (Dautzenberg, 1890; type-locality in Sénégal, West Africa) has been treated as a full species, subspecies of B. truncatus (Audouin) or synonym of the latter. Improved knowledge of B. guernei is desirable as this snail has been implicated as a host in Senegambia for Schistosoma haematobium , the blood parasite causing human urinary schistosomiasis (bilharzia). Snail samples from 18 localities were studied: from The Gambia (3), Sénégal (9), Mali (1), Mauritania (1) and, to represent B. truncatus , Morocco (2) and Egypt (2), Characters investigated were morphological (shell, radula and certain soft parts), biochemical (egg proteins and enzymes from body tissue extracts) and cytological (chromosome number). Multivariate analyses (principal components analysis and canonical variate analysis) were performed on data from the shell and radula. Comparison with the holotype shell of B. guernei confirmed that this taxon is represented by recently collected specimens from Senegambia. Variation in the total material examined did not appear to warrant making any taxonomic subdivision, and thus the taxa guernei and truncatus appear to be conspecific. Further sampling is desirable, particularly to determine the possible value of variation in the enzyme HBDH in defining infraspecific units. Experimental infection showed some individual B. guernei to be a compatible host for 5. haematobium (isolate from Sudan; transmitted locally by B. truncatus ). Snails identified as B. guernei apparently have little importance in the transmission of S. haematobium in the Sénégal river basin at present, but such snails could constitute a health hazard if a compatible form of the parasite were to be introduced into this region of Africa.     

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.     

Uncovering an overlooked radiation: molecular phylogeny and biogeography of Madagascar's endemic river snails (Caenogastropoda: Pachychilidae: Madagasikara gen. nov.)

By analysing key morphological characters (with emphasis on shell, radula and stomach anatomy) and a partial fragment of the mitochondrial 16S rRNA gene (alignment length 860 bp), we examined patterns of diversity and differentiation of a previously overlooked radiation of Madagascan pachychilid freshwater gastropods. These analyses resulted in the discovery of three new species in addition to the two species that were already recognized. The complex nomenclatural and taxonomic implications are discussed and the finding of a viviparous reproductive mode in at least one among otherwise oviparous species is reported. Using a mitochondrial phylogeny that includes all currently accepted pachychilid genera and a strict molecular clock approach, we address the historical biogeography of the Madagascan species with respect to vicariant versus dispersalist biogeographical models. Using two alternative calibrations that were previously suggested for other gastropods, the molecular clock tree suggested that the origin of the Pachychilidae dates back to no more than 50 Mya, whereas the origin of the Madagascan lineage is estimated to date to a period between 15.6–31.5 Mya. These estimates are approximately concurrent with the dating of colonization events in a number of other Madagascan animal taxa. The pachychilid radiation on Madagascar appears not to be older than 3–5 Mya. Thus, although the global patterns of pachychilid distribution have earlier been interpreted to suggest a Gondwanan origin of the family, the present study does not support this postulate. Neither the topology of the molecular phylogeny, nor the timing of events as suggested from a molecular clock were found to be congruent with a vicariance scenario within the framework of Gondwanan fragmentation during the Mesozoic but, instead, imply overseas dispersal during the Cenozoic. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 99 , 867–894.     

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.     

The buccal apparatus with radula of a ceratitic ammonoid from the German Middle Triassic

Complete ammonoid mouth parts including both mandibles and the radula are rare. A newly prepared specimen of Ceratites penndorfi of the late Anisian from the Franconian Muschelkalk reveals one such anaptychus-type jaw apparatus including several more or less clearly recognisable structures such as the shapes and proportions of the inner and outer lamellae of both mandibles. The lower mandible has a short inner and a long outer lamella with an ovoid outline, while the upper mandible has a slightly arched rostrum and the inner lamella carries two wings. The radula, as far as it is preserved, appears to be homodont with oblique, simply conical, monocuspidate teeth. Some other structures are here illustrated, described and interpreted as both mandibles and oesophageal remains. Additional structures of organic origin are preserved such as the radula, but others are difficult to interpret. Some, if not all, of these structures also were body parts of the ceratite, such as perhaps the radular support and/or the oesophagus. All mouthparts, which are preserved in the specimen described herein, are carbonised except for the radula remains, which are phosphatic. The taphonomy of ceratite mouth parts in the Muschelkalk is shortly discussed.     

Larval and juvenile gastropods from a Carboniferous black shale: palaeoecology and implications for the evolution of the Gastropoda

A horizon in the late Visean Ruddle Shale from Arkansas contains the oldest well-preserved gastropod protoconchs known from the Americas. The gastropod fauna consists of a diverse larval shell assemblage and a low diversity assemblage of juvenile gastropods that probably had a benthic life habit. Gastropod larval shells are always isolated, i.e. the gastropods did not complete their life cycle (no metamorphosis) and were unable to become benthic. This was caused by unfavorable environmental conditions on the soft muddy bottom that was probably due to anaerobic to exaerobic conditions. The absence or scarcity of bioturbation caused by invertebrate detritus or sediment feeders in both shale and concretions (formed before compaction) favored preservation of the delicate larval shells. The lack or scarcity of infauna and bioturbation as well as the low diversity of the presumed benthos supports an interpretation of a quasi-anaerobic to exaerobic benthic environment. The superbly preserved larval shells demonstrate that there are more caenogastropod clades present in the late Palaeozoic than suggested previously. Some larval shell types have an openly coiled first whorl followed by a planktotrophic larval shell; openly coiled initial whorls are unknown from modern caenogastropods. The vetigastropods have a smooth protoconch of two whorls clearly demarked from the following whorls - a pattern unknown in modern vetigastropods which have a protoconch of less than one whorl and build no larval shell during their planktonic stage. This could indicate a link between Palaeozoic vetigastropods and the caenogastropods.     

Biology of the Hyolitha

Hyoliths are Paleozoic fossils that have a calcareous exoskeleton consisting of an elongate, usually bilaterally symmetrical cone, a close fitting operculum, and a pair of curved appendages. Their skeletal ultrastructure resembles the crossed-lamellar shell layers of some molluscs. Several specimens from the Ordovician of France and the Cambrian of Antarctica have parts of the gut preserved by infilling matrix, showing that both mouth ad anus were located near the cone aperture. Muscle scars in other hyolith shells indicate that the animal had a series of dorsoventral and longitudinal, or longitudinal and circular muscles, which operated through a hydrostatic skeleton to protract and retract the head, to open and close the operculum, and to move the appendages. Although the shell form and skeletal ultra-structure of hyoliths are of a molluscan type, the muscle insertions suggest that the hyolith cone is not homologous with the dorsal exoskeleton of primitive molluscs. Hyoliths probably constitute a small extinct branch of phylum size, related to the Mollusca and the Sipunculoidea. All three groups may have had common ancestors in the late Precambrian.     

Radula formation in two species of Conoidea (Gastropoda)

The radula is the basic feeding structure in gastropod molluscs and exhibits great morphological diversity that reflects the exceptional anatomical and ecological diversity occurring in these animals. This uniquely molluscan structure is formed in the blind end of the radular sac by specialized cells (membranoblasts and odontoblasts). Secretion type, and the number and shape of the odontoblasts that form each tooth characterize the mode of radula formation. These characteristics vary in different groups of gastropods. Elucidation of this diversity is key to identifying the main patterns of radula formation in Gastropoda. Of particular interest would be a phylogenetically closely related group that is characterized by high variability of the radula. One such group is the large monophyletic superfamily Conoidea, the radula of which is highly variable and may consist of the radular membrane with five teeth per row, or the radular membrane with only two or three teeth per row, or even just two harpoon-like teeth per row without a radular membrane. We studied the radulae of two species of Conoidea (Clavus maestratii Kilburn, Fedosov & Kantor, 2014 [Drilliidae] and, Lophiotoma acuta (Perry, 1811) [Turridae]) using light and electron microscopy. Based on these data and previous studies, we identify the general patterns of the radula formation for all Conoidea: the dorsolateral position of two groups of odontoblasts, uniform size, and shape of odontoblasts, folding of the radula in the radular sac regardless of the radula configuration. The morphology of the subradular epithelium is most likely adaptive to the radula type.     

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)     

Deep water 'dwarfs' and 'giants' among endemic Baikal gastropods

It is suggested that existence of "dwarfs" and "giants" among Baikal gastropods depends on several factors. "Dwarfs" occurs in habitats 1) where sedimentation rate is low and thick silt layer is absent (the small shell without long periostracal nodules does not allow the animal to stay on the soft sediment surface; 2) where there is hard substratum suitable for laying egg capsules; 3) where there is intensive water exchange assuring supply of food (detritus and phytoplankton). Distribution of "giants" capable to lay egg capsules on their own shells is not confined by availability of hard substratum. They dwell on soft sediments because they have large light shell with high whorl allowing them to migrate on silt surface. "Giants" live in places with high sedimentation rates, constant supply of food (including died fish), and higher degree of water mineralization. "Giants" and "dwarfs" inhabit only oxidized substrates. Neither "giants", nor "dwarfs" inhabit the flat bottom of Baikal. "Giants" occur on near-slope parts of the lake, while the "dwarfs" prefer underwater elevations.     

Has Octopus vulgaris a second radula?

Octopus vulgaris drills shelled molluscs and measurements showed the radula to be too large to reach into the depths of the holes made. Small teeth were found on the salivary papilla and its seems likely that these are used for excavating the deeper parts of the holes and penetration of the shell.