THE RELATIONSHIP BETWEEN SHELL INJURIES, SHELL THICKNESS AND HABITAT CHARACTERISTICS OF THE INTERTIDAL SNAIL LITTORINA RUDIS MATON

The hypothesis that risk of shell injury from crab predationand crushing by boulders and stones is important in the selectionof shell thickness in the intertidal gastropod Littorina rudisMaton was tested using data from 24 populations from Britain.Although there was an obvious relationship between the incidenceof shell injuries and both the distribution of crabs and substratummobility, there was no correlation between risk of shell injuryand shell thickness. This is probably accounted for by between-habitatvariations in energy levels which may vary independently ofcrab density and substratum mobility. ^*Present address: Department of Biology, University of Essex,Wivenhoe Park, Colchester C04 3SQ (Received 23 May 1977;     

CORRELATIONS BETWEEN SHAPE, WEIGHT AND THICKNESS OF SHELL IN FOUR POPULATIONS OF VENERUPIS RHOMBOIDES (PENNANT)

Three populations of Venerupis rhomboides in the Plymouth regionand one from the Firth of Clyde show remarkably wide, overlappingranges of variation in shape of shell. Shells of each populationare defined statistically in terms of relationships of shelllength, convexity and mean shell thickness, the latter measuredby a new technique utilizing sections of the shell cut radiallyfrom the umbo. Differences in shell shape are considered againstthe differences in the environments of the four populations,although environmental factors are incompletely known. The results of a transplantation experiment with mature bivalvesare also presented. The survivors showed correlated changesin shell shape and weight in the same direction and with thesame order of magnitude as those found in the distribution ofthe four populations. Detailed examination of the microscopicgrowth line patterns within the shells suggests that the increasedheight/length and convexity/length ratios and increased meanshell thickness were the results of reduced mantle exposureat the shell periphery in conditions of high concentration ofsuspended silt and clay. (Received 12 September 1982;     

THE RATIO OF CALCAREOUS AND ORGANIC SHELL COMPONENTS OF FRESHWATER SPHAERIID CLAMS IN RELATION TO WATER HARDNESS AND TROPHIC CONDITIONS

Shells from 14 populations of sphaeriid clams including Sphaeriumstriatinum, S. simile, Pisidium walkeri, Musculim partumeiumand M. iransversum were analyzed for organic carbon (µgCmg^–1 shell), nitrogen (µg,N mg^–1 shell) andCaCO_s (%CaCO₃ of total clam dry weight). Habitat waters wereassessed for total hardness (expressed as ppm CaCo₃), ppm Ca,ppm Mg, conductivity (µmho) and suspended organic Carbon(µgCl^–1). For all populations, shell organic C andN are positively correlated and there is an inverse relationshipbetween the amounts of shell CaCO₃ and shell organic carbon.Trophic considerations give the best correlation with shelltype at the generic level of consideration since species ofMusculium are found at the opposite end of the trophic scale(eutrophic) from all other populations studied. For S. striatinum,the most extensively studied species, the amount of shell CaCO₃is inversely related to water hardness. The selection of shellsin the Sphaeriidae is discussed in relation to structural-functionalneeds and habitat conditions ¹ Present Address: Department of Biology, Syracuse University,Syracuse, New York 13210, U.S.A. ² Present Address: Department of Zoology, Miami University,Oxford, Ohio 45056, U.S.A. (Received 5 December 1978;     

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;     

INTER AND INTRA-SPECIFIC VARIATION OF THE SHELL OF TWO SPECIES OF THE TROPICAL FRESHWATER SNAIL BELLAMYA (MOLLUSCA: PROSOBRANCHIA)

Shell variation in two species of the tropical freshwater snailBellamya from two different habitats, was studied. The variation of the shell features studied, which includedthe shell width, spire height, aperture height and aperturewidth, were found to be related to shell height. Furthermore,significant difference in the shell shape was found not onlybetween the two species but also between males and females ofeach species. (Received 18 August 1983;     

PREDATION ON THE BIVALVE CHOROMYTILUS MERIDIONALIS (KR.) BY THE GASTROPOD NATICA (TECTONATICA) TECTA ANTON

The effects of a population of the boring gastropod Natica tectaon the bivalve Choromytilus meridionalis were investigated atBailey's Cottage, False Bay, South Africa. In July 1979 theN. tecta density on the mussel bed averaged 69 m^–2 andthe population consisted mainly of reproductively mature individualsbetween 20–33 mm shell width. Laboratory experiments on N. tecta showed that prey size selectionis an increasing function of predator size. The prey size rangetaken by large N. tecta is also greater than that taken by smallindividuals. The position of the borehole on the mussel shellis a function of the way in which the shell is held by the footduring the boring process. Consumption rates measured in thelaboratory showed an increase from approximately 1 kJ per weekin 18 mm N. tecta to 4.5 kJ per week in 28 mm individuals. Populationconsumption in the field was calculated as 663 kJ m^–2month^–1. It was estimated that at this rate the standingcrop of mussels in the pool would be eliminated within 10 months.Field measurements showed significant depletion after 6 months. New spat settlement of mussels occur every 4–6 years.The growth curve shows that after one year the population meansize exceeds 30 mm shell length, which is beyond the prey selectionsize range of small N. tecta. It was concluded that at the timeof a new mussel settlement a niche is provided for the simultaneoussettlement and growth of juvenile N. tecta in high densities.However, within one year the increase in prey size, togetherwith depletion due to over-exploitation, limits population growthand density in N. tecta. (Received 14 March 1980;     

DIFFERENCES IN THE SHELL SHAPE OF LYMNAEA PEREGRA (MULLER) (GASTROPODA: PULMONATA) FROM LOTIC AND LENTIC HABITATS; ENVIRONMENTAL OR GENETIC VARIANCE?

Lymnaea peregra from lotic habitats (running waters) had largeraperture-length/shell-length ratios than contemporaries fromlentic sites (standing waters). This pattern was observed infield samples from two separate years (1985 and 1986). Laboratoryculture experiments suggested that shell shape could be geneticallydetermined in at least some populations. We hypothesize thatlotic snails possess bigger feet, and hence larger aperturesthan lentic individuals as an adaptation to withstand watercurrent ^*Present address Departments of Biology, King's College (KQC),University of London, Campden Hill Road, London W3 7AH, England (Received 14 September 1987;     

Adaptive responses and invasion: the role of plasticity and evolution in snail shell morphology

Invasive species often exhibit either evolved or plastic adaptations in response to spatially varying environmental conditions. We investigated whether evolved or plastic adaptation was driving variation in shell morphology among invasive populations of the New Zealand mud snail (Potamopyrgus antipodarum) in the western United States. We found that invasive populations exhibit considerable shell shape variation and inhabit a variety of flow velocity habitats. We investigated the importance of evolution and plasticity by examining variation in shell morphological traits 1) between the parental and F₁ generations for each population and 2) among populations of the first lab generation (F₁) in a common garden, full‐sib design using Canonical Variate Analyses (CVA). We compared the F₁ generation to the parental lineages and found significant differences in overall shell shape indicating a plastic response. However, when examining differences among the F₁ populations, we found that they maintained among‐population shell shape differences, indicating a genetic response. The F₁ generation exhibited a smaller shell morph more suited to the low‐flow common garden environment within a single generation. Our results suggest that phenotypic plasticity in conjunction with evolution may be driving variation in shell morphology of this widespread invasive snail.     

ALLOZYME VARIATION AMONG POPULATIONS OF BIOMPHALARIA PFEIFFERI (KRAUSS, 1848) (GASTROPODA: PLANORBIDAE) IN CAMEROON

Allozyme electrophoresis was used to study genetic diversityamong the freshwater, hermaphroditic snail Biomphalaria pfeifferiin Cameroon. Four of 19 loci studied were polymorphic. The allozymesfor two loci, aspartate aminotransferase-1 (AAT-1) and isocitratedehydrogenase (IDH), showed distribution patterns related toclimatic zones. AAT-1¹⁰⁰ and IDH¹⁰⁰ occurred predominantly inthe Tropical Climatic Zone, located in the northern half ofthe country. AAT-1¹⁴⁰ and IDH⁹⁰ were common in the EquatorialZone. Intrapopulation variations occurred in 5 of 19 populationsampled. Only one population, polymorphic for AAT-1 locus, waspanmictic. The other 4 populations were not in Hardy-Weinbergequilibrium. The deficiency of heterozygotes in those populationsis unusual for a member of the genus Biomphalaria and may bedue to a predominance of self-fertilization. Other explanationsinclude ecological factors or the presence of segregated sub-populationsthat do not outcross. This latter hypothesis is supported bythe total absence of heterozygotes for PGM alleles. (Received 24 August 1989; accepted 14 September 1989)     

SEXUAL DIMORPHISM OF NUCELLA LAPILLUS (GASTROPODA: MURICIDAE) IN NORTH WALES, UK

Multivariate statistical methods were employed to examine sexualdimorphism in size and shape of Nucella lapillus collected from 16sheltered sites along coasts of Anglesey and the Lleyn Peninsula, NorthWales, UK. Females were significantly larger than males in overallsize; among 12 relative measures of shell shape, two ratios (shellwidth/shell length and aperture length/shell length) were significantlydifferent between males and females, but these differences usuallydecreased with increasing age (shell length). The observed hypoallometricdimorphism could be a result of selection on increased femalefecundity, which may be positively correlated with shell sizein N. lapillus as in other gastropod species. (Received 22 November 1999; accepted 10 April 2000)     

SPATIAL PATTERNS OF SHELL SHAPE OF THREE SPECIES OF CO-EXISTING LITTORINID SNAILS IN NEW SOUTH WALES, AUSTRALIA

The shape and relative weight of the shell have been shown tovary intraspecifically and interspecifically in a number ofspecies of gastropods, including many different littorinids.These differences give rise to different shell forms in differenthabitats. In those species which have non-planktotrophic development,differences in shell form among shores have been usually explainedin terms of natural selection because exposure to waves supposedlyfavours light shells with large apertures, while predation bycrabs on sheltered shores favours elongated, thick shells withsmaller apertures. Differences in shell shape among speciesfound at different heights on the shore have been explainedin terms of resistance to desiccation and temperature. Suchvariables would tend to act on a relatively broad-scale, i.e.causing differences among heights on a shore or among shores.Rates of growth, which might vary at much smaller scales withina shore, have also been shown to affect the shapes of many shells. In this study, the shape and relative weight of shells of threespecies of co-existing littorinids (Littorina unifasciata, Bembiciumnanum and Nodilittorina pyramidalis) were measured. These speciesall haveplanktotrophic development and they are found on manyshores where there is no evidence that they are preyed uponby crabs. Before explanations of shell shape are proposed, itis necessary that patterns of variation, within different partsof ashore and among different shores are clearly documented.These patterns were measured at a number of different spatialscales within and among replicate shores with different amountsof wave exposure. Large and small specimens were included toallow intraspecific comparisons among snails of different sizesfound at different heights on the shore. The results showedsignificant differences among shores in shape and relative weightof shells, but these differences could not be explained by exposureto waves. In addition, snails of different sizes and differentspecies did not show the same patterns although they were collectedfrom the same sites. Importantly, the shell shape of Liuorinaunifasciata varied significantly among sites at approximatelythe same height within a shore. These differences could notbe clearly correlated with density, mean size nor exposure towaves. The only consistent pattern was a decrease in relativeaperture size in specimens living higher on the shore. Modelsthat have commonly been proposed to explain shape and relativeweight of shells in other species of gastropods are not adequateto explain the small- and large-scale variation of the measurementsdescribed here. It is proposed that any selective advantageof shell morphology and the effects of any variables on thedevelopment of shell morphology in these species can only beidentified after appropriately designed and replicated fieldexperiments. (Received 4 March 1994; accepted 13 September 1994)     

SOME ENZYMES RELATING TO THE METABOLISM OF STARCH IN A SEA-LETTUCE

The occurrence of a phosphorylase in the extracts of a greenseaweed, Ulva pertusa was demonstrated. In addition, a sugarphosphatase different from non-specific acid phosphatase andan amylase, which are involved in the starch degradation, werealso detected in this sea-lettuce. ¹Contributions from the Shimoda Marine Biological Station, No.147.     

TRANSFER EXPERIMENT SUGGESTS ENVIRONMENTAL EFFECTS ON THE RADULA OF LITTORARIA FLAVA (GASTROPODA: LITTORINIDAE)

Phenotypic variation in radulae has been studied in severallittorinid species because of this organ's intrinsic relationshipwith diet and, consequently, with the environment. In this work,we compared the radulae of the Brazilian species Littorariaflava found in mangroves and on rocky shores. Individuals ofL. flava showed marked differences in the shape of the cuspsamong samples from rocky shore and mangrove. In a transfer experiment,the shape of the radula changed within 40 days. A differentresponse was observed in individuals transferred to mangrove,where two different phenotypes were found, suggesting eitherintrapopulational variation in the responses to change of environmentalconditions, or that some snails showed a slow reaction to theenvironmental changes. The alterations could be attributed toecophenotypic plasticity. Analysis of variance showed that thelength of the radula in L. flava was strongly influenced bythe substrate (F_6,22=17.13, P<0.000), but apparently notby the transfer experiment. (Received 3 May 2005; accepted 18 July 2005)     

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)     

A Population Study to Aid the Selection of Improved Dried Pea (Pisum sativum) Crop Plants

The effect of genotype and plant density, over the range from100 to 277 plants m^–2, on plant to plant variation inprecision sown microplots has been assessed for three ‘leafless’(afafstst) pea (Pisum sativum) lines. This range of plantingdensities did not significantly affect the total above groundbiological yield per unit area of two of the genotypes (BS5and BS4) whereas the biological yield of the third (BS151) declinedat densities above 156 plants m^–2. The differences weredue to changes in seed yield. The effect of planting densityon the variation between plants for biological yield withinthe microplots differed between the genotypes. The distributionpattern of BS4 and BS5 changed from normal to skewed with increaseddensity, while the distribution for BS151 remained skewed atall planting densities. The differences between the three genotypes in the proportionof biological yield partitioned into seed yield (harvest index)on a unit area basis was due almost entirely to the differencesin structure of the plant populations. The maximum level ofpartitioning by individual plants was similar for all threelines. The difference between this maximum for an individualand the crop harvest index therefore represents the area forimprovement of crop harvest index through breeding. It is suggestedthat improvements in dried pea yields will come, therefore,by selecting plants which form more uniform populations withregard to plant size and to the proportion of plant biomasspartitioned into seed (plant harvest index). Pisum sativum, leafless pea, population, genetic variation, distribution patterns, harvest index     

EFFECTS OF WOUND HEALING ON ZINC DISTRIBUTION AND ALKALINE PHOSPHATASE ACTIVITY OF HELIX ASPERSA (GASTROPODA: PULMONATA)

Helix aspersa were loaded with ⁶⁵zinc and after 2 and 5 daysstarvation there was no significant reduction in the zinc concentrationof the digestive gland and shell. Wounding of the optic tentacle,foot tissue and partial shell removal of H. aspersa resultedin deposition of ⁶⁵zinc in the wound area. Alkaline phosphataseshowed no increase in concentration at the sites of wounding. (Received 15 October 1985;     

PRELIMINARY INVESTIGATIONS OF THE ECOLOGY OF THE INTERTIDAL PREDATORY GASTROPOD LEPSIELLA VINOSA (LAMARCK) (GASTROPODA MURICIDAE)

Lepsiella vinosa\ (Lamarck), a common intertidal predatory gastropod,was studied in two zones on an exposed rocky shore in Victoria(Australia). Those from the Mussel Zone in the mid-shore primarilyate one species of mussel and had a faster rate of energy consumptionthan those from the Littorinid Zone in the high-shore, whichmainly ate one species of littorinid gastropod. Shell length,growth rate and size-specific body weight of L. vinosa wereall significantly greater in the Mussel Zone. Egg capsules ofL. vinosa from the Mussel Zone contained significantly moreeggs and more potential hatchlings per capsule than those fromthe Littorinid Zone although there were no significant differencesin the sizes of eggs or hatchlings between zones. These differencesbetween the two zones were correlated to the greater consumptionof energy by L. vinosa in the Mussel Zone and were discussedin relation to the absence of dispersive larvae in this species ¹Present address: Department of Zoology, Monash University,Clayton, Victoria, 3168, Australia ²Present address: School of Biological Sciences, Carslaw Building,F07, University of Sydney, N.S.W., 2006, Australia. Addresseefor reprints. (Received 2 June 1987;     

ALLOZYME COMPARISONS AND PLOIDY LEVELS AMONG SPECIES OF THE BULINUS TRUNCATUS/TROPICUS COMPLEX (GASTROPODA: PLANORBIDAE) IN CAMEROON

Allozyme electrophoresis, ploidy level, and, to a lesser extent,anatomy and shell morphology were used to characterize snailsof the Bulinus truncatus/tropicus complex in Cameroon. Starchgel electrophoresis revealed no intrapopulation variation within20 populations examined for 14 allozyme loci in 12 enzyme systems.Interpopulation differences in allozyme patterns were used todivide the populations studied into five groups. Each groupis distinguished by a unique combination of zymograms for hydroxybutyratedehydrogenase and isocitrate dehydrogenase. Four of these groupsare tetraploids and are identified as B. truncatus. Populationsof the most commonly found B. truncatus group are generallyaphallic, are frequently associated with transmission of S.haematobium, and generally occur at altitudes below 800 m abovesea-level. Populations of B. truncatus belonging to the otherthree groups are usually euphallic, not associated with S. haematobiumtransmision, and generally occur at altitudes above 1000 m.These four groups may represent distinct biotypes of B. truncatus.Populations belonging to the fifth group are diploid, euphallic,not associated with S. haematobium transmission, and occur inhabitats above 1000 m. This is the first confirmed report ofdiploid B. truncatus/tropicus-complex snails from this regionof Africa. They are tentatively identified as B. natalensis,a species previously thought to be limited to East and SouthernAfrica. (Received 31 October 1988; accepted 20 February 1989)     

Shell shape variation in the Nassariid Buccinanops globulosus in northern Patagonia

Morphological variation among natural populations is a phenomenon commonly observed in marine invertebrates and well studied, particularly, in shelled gastropods. The nassariid Buccinanops globulosus is interesting to study shell shape variation because it exhibits strong interpopulation differences in life history features, including maximum size, fecundity and growth rate. In this study, we examined the pattern of variation in size and shell shape among populations and between sexes of B. globulosus (Bahía San Antonio 40°29′S 63°01′W, Playa Villarino 40°45′S 64°40′W and Bahía Nueva 42°46′S 65°02′W). In particular, we used geometric morphometric techniques to test: (1) whether the two components of shell morphology (size and shape) are independent and (2) whether shape differences between sexes are consistently found among populations, regardless of their body sizes. Our results show shell shape variation between the populations of B. globulosus of northern Patagonia. Intra-specific shell shape variation is affected by body size, indicating allometry. Regardless of the size differences, individuals from Playa Villarino have high-spired shells, and shorter apertures and wider columellar area than individuals from the other populations. Also, sex-related shape differences were consistently found at each population, thus suggesting a common sexual dimorphism in shell morphology for this species. The functional significance of the variability found is discussed in terms of the flexibility of developmental programmes for morphology as well as the evolution of phenotypic plasticity.     

PHOSPHOGLUCOMUTASE ALLOZYME EVIDENCE FOR AN OUTCROSSING MODE OF REPRODUCTION IN THE HERMAPHRODITIC BROODING BIVALVE MYSELLA TUMIDA (GALEOMMATACEA)

Allozyme variation of a Mysella tumida population in PatriciaBay, B.C., Canada was investigated at the phosphoglucomutase(PGM) locus. Seven alleles were detected. The observed frequencyof allelic combinations did not differ significantly from randommating expectations (0.25 < P < 0.50), however, heterozygositylevels were slightly greater than expected (Selander's D = 0.022).These results, together with previous work on the M. tumidareproductive cycle, indicate that natural populations of thisbivalve rarely, if ever, self-fertilize. ^*Present Address: Friday Harbor Laboratories, University ofWashington, Friday Harbor, WA 98250, U.S.A. ^**Present Address: Museum of Zoology, University of Michigan,Ann Arbor, M1 48109, U.S.A. (Received 30 December 1986;