CONTRIBUTION OF GENETIC AND ENVIRONMENTAL FACTORS TO SHELL SHAPE VARIATION IN THE LOTIC SNAIL SEMISULCOSPIRA REINIANA (PROSOBRANCHIA: PLEUROCERIDAE)

The genetic and environmental factors affecting shell shapein the freshwater snail Semisulcospira reiniana at Takahashiin Kyoto, central Japan, were studied by means of a rearingexperiment and field observations. Shell shape was characterizedby three parameters; W (whorl expansion rate), T (whorl translationrate), and S (roundness of generating curve). Estimated heritabilitieswere low in all three parameters and the largest component ofthe great shell variation in the Takahashi population was environmentalvariance, suggesting that the main source of shell variationwas phenotypic modulation, in response to the external environment.In the field, snails that were active in fast currents had largerW and smaller T, that is, a larger body whorl and a lower spire,than snails in slow currents. Substratum conditions relatedto T in resting periods although its cause was unclear. Mechanismsare suggested which in the absence of selection of genetic variationcould cause and maintain shell variation in S. reiniana in differentmicrohabitats. (Received 8 March 1996; accepted 3 November 1997)     

LIFE CYCLE AND HABITAT SHIFT OF THE TROCHID SNAIL DILOMA SUAVIS (PHILIPPI) WITHIN INTERTIDAL MUSSEL ZONES

The life cycle of the trochid snail Diloma suavis (Philippi, 1849),was studied on an intertidal rocky shore at Shirahama, Wakayama Prefecture,where two mytilid bivalves, Septifer virgatus (Wiegmann) andHormomya mutabilis (Gould), formed vertically contiguous musselbeds in the upper-middle and lower zones, respectively. At lowtide in April, the snail density increased with decreasing shoreheight and was greatest at the middle level of the H. mutabilisbed. Then, the density decreased towards the lower littoralfringe. Newly settled juveniles smaller than 2 mm in shell heightappeared abundantly in late summer and autumn within algal turfon the lower shore. As snails grew larger than 2 mm, they appearedwithin the gaps of the H. mutabilis bed and the S. virgatusbed. They increased in size monotonically towards the next summer,but rate of growth in shell height tended to be great in autumnand small in winter. Seasonal change in the density of snailsfound within the gaps of the mussel beds was remarkable during athree year period, increasing from autumn to winter and then decreasingtowards next summer. Reproduction occurred in summer, and adultsnails disappeared by September. It is thus suggested that this specieshas a one year lifespan and shows a habitat shift from algal turfto the gaps of the mussel beds with growth. (Received 12 October 1998; accepted 2 March 1999)     

INTRASPECIFIC AND INTERSPECIFIC VARIATION IN SHELL MORPHOLOGY OF BIOMPHALARIA PFEIFFERI (KRAUSS, 1848) AND HELISOMA DURYI (WETHERBY, 1879) FROM MOSHI, TANZANIA, ANALYSED BY MORPHOMETRIC STUDIES

Helisoma duryi, a planorbid snail species, which is a potentialbiological control agent of the intermediate hosts of schistosomiasis,was introduced in an irrigation scheme (T.P.C.) in northernTanzania in 1972. The species has been present at T.P.C. since,and during a snail survey in January 1981, a great variationin the shell morphology of H. duryi, ranging from typical H.duryi forms to forms resembling Biomphalaria pfeifferi, theintermediate host of Schistosoma mansoni in the area, was observed.The Biomphalaria- Wkc forms of H. duryi could constitute anentire population but also occasionally occurred in populationsof typical H. duryi. This study was undertaken to determine whether simple shellmeasurements, or ratios between some of these could be usefulin discriminating between H. duryi and B. pfeifferi. Followingparameters were considered: shell height, shell diameter, diameterof the umbilicus, ratio between shell height and diameter ofthe umbilicus, and the number of whorls. (Received 29 July 1983;     

DISTRIBUTION AND ABUNDANCE OF HYDROBIA ULVAE (PENNANT) IN THE LOWER MEDWAY ESTUARY, KENT

Measurements of densities and distribution of populations ofH. ulvae were made in the lower Medway estuaryduring the period1969–1975, together with measurements of salinity, substrateparticle size and macrophyte distribution. Results indicatedthat sheltered siteswere more favourable to the snail as shownby higher population density and greater shell size. ^*Current address: Southern Water Authority, Maidstone, Kent (Received 20 May 1979;     

DREPANOTREMA NORDESTENSE (LUCENA, 1953) (PULMONATA: PLANORBIDAE): A NEW RECORD FOR THE BOLIVIAN ALTIPLANO AND ITS DIFFERENTIATION BY MORPHOLOGY AND ENZYMES

In 1995, a small planorbid snail new for the Northern Bolivian Altiplanowas discovered in several temporary ditches. Conchological andanatomical characteristics of this snail are similar to thoseof Drepanotrema nordestense (Lucena 1953), a tropical planorbid snailreported from the northeastern region of Brazil. A comparative studywas carried out with the closely related species D. aeruginosumfrom Puerto Rico and Guadeloupe using morphological (shell andanatomy of the reproductive system) and genetic (allozyme) markers.This study confirmed a clear distinction between the two species.Several malacological surveys have been carried out in the NorthernBolivian Altiplano since the first field trip by D'Orbigny in1835, the last one in 1975. That D. nordestense and relatedspecies were not reported in these earlier surveys may indicatea relatively recent introduction. (Received 6 March 2000; accepted 1 June 2000)     

AGE DETERMINATION OF ARIANTA ARBUSTORUM (L.) (PULMONATA) BASED ON GROWTH BREAKS AND INNER LAYERS

The pulmonate snail Arianta arbustorum from the Swiss Alps wasaged using thin sections of the shell margins cut from markedindividuals. Shell layers at the apertural lip and growth breaksin the juvenile can give a reliable estimate of age. (Received 16 May 1986;     

Bioluminescent signals spatially amplified by wavelength-specific diffusion through the shell of a marine snail

Some living organisms produce visible light (bioluminescence) for intra- or interspecific visual communication. Here, we describe a remarkable bioluminescent adaptation in the marine snail Hinea brasiliana. This species produces a luminous display in response to mechanical stimulation caused by encounters with other motile organisms. The light is produced from discrete areas on the snail's body beneath the snail's shell, and must thus overcome this structural barrier to be viewed by an external receiver. The diffusion and transmission efficiency of the shell is greater than a commercial diffuser reference material. Most strikingly, the shell, although opaque and pigmented, selectively diffuses the blue-green wavelength of the species bioluminescence. This diffusion generates a luminous display that is enlarged relative to the original light source. This unusual shell thus allows spatially amplified outward transmission of light communication signals from the snail, while allowing the animal to remain safely inside its hard protective shell.     

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)     

ROLE OF FOOD IN MASS CULTIVATION OF THE FRESHWATER SNAIL INDOPLANORBIS EXUSTUS, VECTOR OF ANIMAL SCHISTOSOMIASIS

For the mass cultivation of the tropical freshwater snail, Indoplanorbisexustus, primary intermediate host of animal schistosomiasis,three foods, spinach, sheep's liver and a synthetic food (M/sHindustan Lever's rat food) were evaluated. The effect of thesefoods on growth, sexual maturity, fecundity and mortality isconsidered in this study. It is concluded that synthetic foodis ideal for breeding and sheep's liver for the maintenanceof mass cultures of this snail. revised 27 August 1985;     

Variation in adult shell morphology and life-history traits in the land snail Oreohelix cooperi in relation to biotic and abiotic factors

In this study, we explore factors influencing variation in adultshell morphology and life-history characteristics (offspringsize and number) in the ovoviviparous land snail, Oreohelixcooperi, from the Black Hills, South Dakota and Wyoming, USA.To date, no study of shell morphology and life-history characteristicsin ovoviviparous land snails has included a combination of datasetsincluding genetic data, life-history traits, shell morphologyand multiple environmental factors. We report that differencesin shell size among populations are strongly related to meanannual temperature (and the highly correlated variable elevation)and population density (measured as shell density). In addition,shell size varies among populations, despite an apparent lackof population genetic differentiation. Common factors thoughtto influence adult shell size, like precipitation and calciumlevels, do not have a significant effect in this study. Adultsize strongly influences per-clutch reproductive output, withlarger snails having larger and more offspring. As mean annualtemperature and shell density affect adult shell size, theyalso indirectly affect per-clutch output. The results suggestthat a large portion of the life-history variation in O. cooperiis environmentally induced, as has been found in oviparous landsnails and brooding freshwater bivalves. (Received 20 July 2006; accepted 10 January 2007)     

ALTITUDINAL VARIATION AND ADAPTIVE ZONES IN ARIANTA ARBUSTORUM: A NEW LOOK AT A WIDESPREAD SPECIES

Arianta arbustorum is a European land snail, occurring in bothlowland and mountainous regions. Its shell is usually globular,which makes the species aberrant among the Campylaeinae in shellshape, habitat, and wide, not exclusively montane range. Withincreasing altitudes, the shells usually decrease in size andbecome more conical. However, in some mountain stocks thereare populations with depressed shells, provided with an openumbilicus. This form (subspecies) is only known from formerPleistocene refugia. It is hypothesized that in the Pliocene, the primary montaneA. arbustorum, with a depressed shell, entered an additional,adaptive zone, while conquering damp lowland habitats. Thisresulted in an enormous range extension. In the new habitata different (globular) shell shape evolved and the species becamepolytypic. The subsequent glaciations wiped out most of theoriginal montane A. arbustorum with depressed shells. Wherethis had occurred, post-Pleistocene re-colonizing conspecificsnails with globular shells, found vacant territory and coulddevelop the dines mentioned before. The invading populationsdid not regain the original, montane, depressed shell shape.This shape was only retained in a few refugia, where the populationssurvived amidst the glaciers. (Received 9 February 1990; accepted 31 May 1990)     

Boring of Shell by Caobangia in Freshwater Snails of Southeast Asia

Giard (1893) described Caobangia billeti, a freshwater sabellidpolychaete, based on material collected in what is now northernNorth Vietnam. Up to the present time it had not been re-collectedand Giard's observations that it is hermaphroditic, that itseggs undergo internal development, and that, rather than havinga posterior terminal anal opening, the anus opens far forward,have been questioned from time to time. Recently collected material, as well as older material in thecollections of the Division of Molluscs, Museum of Natural History,Smithsonian Institution, has allowed the confirmation ofGiard'sfindings and the observation of the burrows of Caobangia in22 species of molluscs from fast-moving streams in Ceylon, northernIndia, Assam, Burma, Thailand, Laos, Java, Sabah, and variousof the Philippine Islands. Burrowing in snail shells appears to take place subsequent tolarval metamorphosis as youngindividuals grow posteriorly froma capsule-like structure through the periostracum and into thecalcium carbonate of the snail shell. There appears to be nodifferentiated area of the worm which can be implicated as thesite of acidic secretions, and the setae of the worms seem notto be unusually worn, as would be the case if they were usedto wear away the shell material.     

Excavation of Boreholes by the Gastropod, Urosalpinx: An Analysis by Light and Scanning Electron Microscopy

Penetration of shell by the muricid gastropod, Urosalpinx cinereafollyensis, is accomplished by successive alternating periodsof (a) chemical activity by the accessory boring organ (ABO), and (b) rasping by the radula. This paper reports on the functionsof the radula and of the ABO in producing the characteristicgeometry of the borehole, andon the effects of radular teethand of the ABO secretion on the microscopic anatomy of the surfaceof the borehole during the process of shell-boring. Radulae of U. c. follyensis and the surfaces of incomplete boreholesin the shell of Crassoslrea virginica, Mytilus edulis, and Myawere examined by means of light and scanning electron microscopy.Hardness tests of radular teeth andshell of prey demonstratedthat marginal teeth are harder than rachidian teeth, and thatthe range of hardness of rachidianteeth overlaps that of thethree species of shell. Rasping is carried out by two, occasionallythree, of the five rachidiancusps. Rasping patterns are shallowand asymmetric. Rachidian teeth are worn to the base with use;marginal teeth wear onlyslightly as they are employed mainlyin feeding. The distance between the tips of rachidian cuspscorresponds with the interval between the parallel cusp tracesrasped by them in shell. During each rasping period, snailsscrape off about 1/10 to 1/5 of the surface of the chemicallytreated area of the bottom of the borehole. Dissolution of shell is accomplished by secretion from the secretorydisk of the ABO. With each application of the ABO,most or allof the radular marks of the previous rasping period are erasedby solution of a thin layer of shell. The pattern of etchingis specific for each of the species of shell studied. In oysterand mussel shell, initial solubilization occurs through theorganic, non-mineralized, prism sheaths, exposing prismaticforms shown by other workers to be distinctive for these species,and then proceeds into the organic-calcareous structure of individualprisms. Etching of Mya shell revealed no fundamental prismaticform. Shell-penetration includes dissolution of both organiccomplexes and CaCO₃ crystals. Shell-boring by this snail is principally a chemical process,and the geometry of the borehole is generally a reflection ofthe morphology of the ABO.     

Trade-offs between force and fit: extreme morphologies associated with feeding behavior in carabid beetles

We explored how functional trade-offs in resource handling strategies are associated with the divergent morphology of predators. The malacophagous carabid Damaster blaptoides shows two extreme morphologies in the forebody; there is an elongate small-headed type and a stout large-headed type. A feeding experiment showed that the small-headed type obtained a high feeding performance on snails with a thick shell and a large aperture by penetrating the shell with its head. In contrast, the large-headed type showed a high feeding performance on snails that had a thin shell and a small aperture, and they ate these prey by crushing the shell. The large-headed, strong-jawed beetles are efficient at shell crushing but are ineffective at shell entry; the large mandibles and musculature that allow for shell crushing make the beetle's head too wide to penetrate shell apertures. On the other hand, small-headed, weak-jawed beetles crush poorly but can reach into shells for direct predation on snail bodies. These findings are hypothesized to be functional trade-offs between force and fit due to morphological constraints. This trade-off would be a primary mechanism affecting both resource handling ability in animals and phenotypic diversity in predators and prey.     

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.     

Functional Morphology of Perihelia conradi Relative to Shell-Penetration

The pholad, Penitella conradi, is found along the Californiacoast in the calcareous shell of the abalone, Haliotis rufescens.These pholads penetrate the abalone shell, and when they breakthrough the inside of the shell they cease boring, secrete acallum, and then become sexually mature. The normal adult isa stenomorphic form,defined by Bartsch as an animal whose sexualmaturity is induced by over-crowding or insufficient substratumin which to bore. In the case of P. conradi, sexual maturityis always induced by the spatial limits of the substratum, thatis, the relatively thin abalone shell. The role of mechanical abrasion by the valves of P. conradiis minor. Experiments indicate that the teeth of P. conradiare worn at a greater rate than the polished shell of the abalone. The boring process in P. conradi proceeds mainly by chemicaldissolution of the calcareous substrate. The pathway of thesolvents is unknown. It may be through the organic matrix, orthe solvent may react directly with the crystals. Mechanicalabrasion helps to remove loosened crystals and/or organic matrixwhich are then carried to the exterior by the ciliary currentsflowing in through the pedal gape and out through the exhalentsiphon.     

GENETIC AND MORPHOLOGICAL VARIATION OF POPULATIONS BELONGING TO THE BULINUS TRUNCATUS/TROPICUS COMPLEX (GASTROPODA; PLANORBIDAE) IN SOUTH WESTERN ZIMBABWE

Aquatic snails from south western Zimbabwe belonging to theBulinus trunscatus/tropicus complex vary widely in shell formsuggesting that more than one taxon could be present. This possibilitywas investigated by making observations on snail samples from13 populations from the Plumtree area, in respect of allozymevariation (5 polymorphic loci), shell morphology (9 variables),copulatory organ and chromosome number. Comparative data wereobtained from snails from north western Zimbabwe identifieddefinitely as B. tropicus. Analysis of the genetic structurerevealed a high degree of polymorphism (P) ranging from 0.29–0.80among populations from Plumtree and expected heterozygosity(H_e) from 0.02–0.22. No enzymatic diagnostic loci werefound which could differentiate the different morphs or populationsand discriminant function analysis on the morphological datashowed an overlap of morphs among populations. Snails analyzedfor chromosome number were all diploid (2n = 36). Snails exposedto Schistosoma haematobium mira-cidia were all refractory. Thisinformation supports the view of a single species, B. tropicus,which is differentiated due to migration barriers and whereenvironmental variables might be implicated in the morphometricdivergence. (Received 31 July 1995; accepted 15 January 1998)     

THE EARLY LIFE-CYCLE STAGES OF HYDROBIA VENTROSA AND HYDROBIA NEGLECTA WITH OBSERVATIONS ON POTAMOPYRGUS JENKINSI

The egg mass of Hydrobia neglecta contains a single egg, whilethat of H. ventrosa contains up to three eggs. At hatching H.neglecta has a significantly greater shell length than H. ventrosaand the late embryos and young snails of the two species canbe separated according to the pigmentation of the head region.The young snails can also be separated on the basis of surfaceornamentation of the shell and comparisons with the closelyrelated H. ulvae and Potamopyrgus jenkinsi show how the basicpattern of shell sculpturing differs in the four species. Thesedifferences are discussed in relation to the mode of development. (Received 14 February 1980;     

SHELL STRENGTH OF COLOUR MORPHS OF THE MANGROVE SNAIL LITTORARIA PALLESCENS

The load required to break shells of the snail Littor-aria pallescens(Philippi) (Gastropoda: Littorinidae) has been established.Mass increases more than proportionally with size. Size forsize, relatively heavy shells are thicker than relatively lightones, and the thicker shells are stronger. Three samples fromdifferent localities differ significantly in strength. Littorariapallescens is polymorphic for shell colour. It is shown thatin two of the three samples, both from Papua New Guinea, individualsof the dark morph are significantly heavier and stronger thanthose of the yellow morph, while there is no difference betweenmorphs in the third sample, from Thailand. In the sample withthe most robust shells the dark individuals are stronger, massfor mass, than yellows. Thus, colour morph is sometimes, butnot always, associated with differences in shell characteristicswhich are likely to affect fitness. Where there is a difference,the dark morph is estimated to be on average about 22% strongerthan the yellow morph. The difference between morphs is smallcompared with the large variation in shell characters associatedwith size. (Received 20 May 1992; accepted 22 June 1992)     

TWO NEW SPECIES OF THE GENUS ISLAMIA RADOMAN, 1973 FROM ITALIAN ISLANDS (PROSOBRANCHIA, HYDROBIIDAE)

Two new Italian species of the genus Islamia (Prosobranchia:Hydrobiidae), one living in eastern Sicily (I. cianensis), andone on Elba Island (Tuscan Archipelago, Italy) (I. gaiteri)are described. The two species are distinguished on the basisof shell and anatomical characters, mainly those of the malegenitalia. I. cianesis n. sp. is characterized by a valvatoidshell and a penial lobe with internal band of glandular tissuenot distinct in its lower portion from the penis body but bulgingapically as a small knob. I. gaiteri n. sp. is characterizedby a planispiral shell and a small lateral penial lobe withoutinternal glandular tissue. (Received 20 December 1993; accepted 27 June 1994)