INTRA-POPULATION VARIATION OF THE SHELL OF LITTORINA RUDIS (MATON) (MOLLUSCA: PROSOBRANCHIA)

Shell variation within a single population of Litlorina rudis(Maton), collected near the Biological Station at Trondheim,Norway, was studied. The variation of the shell features studied, which includedthe shape of the basal part of the outer lip, and the relativeheight of the spire, width of the shell and of the aperture,were found to be at least partially related to shell height.Furthermore, the shell shape in the population investigated,besides varying with age, also varies due to shell damage. Nosignificant differences were found between the shapes of theshells of males and females. The shell characteristics of different populations of L. rudisalso vary greatly, one of the causes being differences in theenvironmental conditions. When studying this variation betweendifferent populations, however, it must be emphasized that onlyundamaged shells of individuals of about the same age shouldbe compared, in order to obviate the effects of the variabilitydue to age differences and shell damage within each individualpopulation. ^*Contribution from Trondhjem Biological Station no. 199 (Received 18 February 1980;     

A COMPARATIVE STUDY OF THE MORPHOLOGY OF THE PULMONATE SNAIL PSEUDOTACHEA LITTURATA (PFEIFFER) AND OTHER SPECIES OF PSEUDOTACHEA, IBERUS AND CEPAEA

A population of Pseudotachea litturata (Pfeiffer, 1851) fromTarifa (Càdiz, Spain) has been studied. The morphologicalresults are compared with those from P. splendida, Iberus gualtierianus,I. alonensis, I. marmoralus, I. guiraoanus and four speciesof the genus Cepaea using, as an exploratory method, the Wagnerparsimony procedure and 18 characters of the shell, genitalsystem and karyotype have been analysed. According to this methodit seems that the taxonomical position of P. litturata in thegenus Pseudotachea is confirmed, and agrees with the phylogeneticalrelationships in this group of species. The genus Cepaea seemsto be well established, although two species groups can be distinguished:C. nemoralis—C. hortensis and C. syluatica—C. vindobonensis.These differ mainly in chromosome number, diverticulum lengthand degree of shell polymorphism. Although the present resultsdo not allow us to clarify the current taxonomical problemswithin the genus Iberus, the species studied seem to belongto a natural group (Received 15 September 1987; accepted 1 January 1988)     

GENETIC VARIATION IN SIX SPECIES OF SACOGLOSSAN OPISTHOBRANCHS

Genetic variation in allozymes of six species of Saco-glossa(Mollusca, Opisthobranchia) from Denmark was studied. Nine enzymesrepresenting 11 loci gave reactions that could be used to calculateallele frequencies. Elysia viridis, Limapontia capitata, L.de-pressa, Ercolania nigra, Alderia modesta and Calliopaea oophagaall showed very great variation both within and between species.Levels of polymorphism varied between 0.5 and 0.8, and levelsof observed heterozygosity were between 0.18 and 0.42. Interspecificidentity of alleles in polymorphic loci was only found betweenthe two species of Limapontia. Even in these two species geneticidentity (I) was very low (0.07) for congeneric species. Theresults are discussed in relation to habitat, fertility, developmenttype and geographic range of the species. (Received 28 February 1990; accepted 26 July 1990)     

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)     

GEOGRAPHICAL VARIATION OF SHELL MORPHOLOGY IN CYPRAEA ANNULUS (GASTROPODA: CYPRAEIDAE)

Macro- and microgeographic variation of shell morphology inCypraea annulus was studied. Surveys of phenotypic variationin 19 populations from the Indian and West Pacific Oceans revealeda significant positive relationship between average seawatertemperature and callus thickness. In contrast, juvenile shellsize after maturation had no significant relationship with seawatertemperature. Research on within-population variation in OkinawaIsland indicated that shell morphology is correlated with someecological factors such as temperature and algal abundance.Both juvenile shell size and callus thickness decreased withincreasing density of individuals. These microgeographical patternssuggest that ecophenotypic variation, rather than genetic differentiation,causes the latitudinal clines in shell morphology that havebeen reported in several cypraeid species. (Received 18 November 2004; accepted 8 April 2005)     

THE AMINO ACID COMPOSITION OF FRESHWATER MOLLUSC SHELLS IN RELATION TO PHYLOGENY AND ENVIRONMENT

Quantitative thin layer chromatography of structural amino acidswas standardized for the study of shell proteins of 13 speciesof freshwater mollusc. There was slight intraspecific variationbetween individuals, particularly for tyrosine concentrations.No external periostracum was chemically identifiable in theshell of Lymnaea peregra (Müller) and this species showedsome amino acid heterogeneity in different parts of the shell.Despite these intraspecific variations, analysis of variancesuggested interspecific variation in concentrations for almostall the amino acids tudied. Cluster and principal componentsanalysis indicated that the amino acid composition did reflectphylogenetic affinity but that environmental factors were probablymore-important. (Received 5 August 1982; revised 5 August 1982;     

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)     

PREDATION ON YOUNG PECTEN MAXIMUS (L.) (BIVALVIA), BY THE ANEMONE ANTHOPLEURA BALLII (COCKS)

On the S.W. coast of Ireland at Lough Hyne (Ine) to the southof the rapids, an area of strong water flow, the anemone Anthopleuraballii (Cocks) was found preying on young escallops, Pectenmaximus (L.), in situ and under controlled conditions. Completedead shells were commonly found beside or on the oral surfaceof anemones. Anemones fed under controlled conditions digestedyoung escallops (8 - 17 mm shell height) within 7 - 15 h andthe ejected shell was distributed the same way. An undisturbedanemone accepted and devoured three spat presented to it insuccession. This species of anemone is locally common and isoften found in areas where escallops are encountered, and maythere cause significant mortalities. (Received 16 October 1982;     

ALLOZYME COMPARISON OF LITTORINA SPECIES FROM THE NORTHWESTERN PACIFIC

Three unknown Littorina species were revealed by allozyme studyof snails from the northwestern Pacific. One of them was foundin Kasatka Bay (Pacific coast of Iturup Island, south KurileIslands) together with L. sitkana, which it resembled in anatomicalfeatures, but the genetic distance between these species wasD=0.622. The second one was found in Tauiskaya Inlet (northernpart of Okhotsk Sea) and was again similar to L. sitkana inmorphological and anatomical features, but D=1.160. The thirdspecies was found in Egvekinot Inlet (Anadyrskiy Gulf, BeringSea) in sympatry with L. natica, to which was close genetically(D=0.256) although it differed in shell shape. (Received 4 April 2005; accepted 26 August 2005)     

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;     

A NEW OVOVIVIPAROUS SPECIES OF TECTARIUS (GASTROPODA: LITTORINIDAE) FROM NIUE, SOUTH PACIFIC, WITH A MOLECULAR PHYLOGENY OF THE GENUS

A new ovoviviparous littorinid gastropod, Tectarius (Echininiopsis)niuensis, from Niue, west of the Cook Islands, is described.This is distinguished from the only other ovoviviparous memberof the genus, T. (E.) viviparus (Rosewater, 1982) from the MarianaIslands, here redescribed in detail. The new species is remarkabletor its high-level habitat in the littoral fringe on wave-exposedkarstic limestone cliffs, for its variation in shell shape accordingto tidal level, shell colour polymorphism, calcified operculumand penis with a single mamilliform penial gland. In a cladisticanalysis of morphological characters, including single representativesof each of the three other subgenera of Tectarius (Tectarius,Echininus, Tectininus), these two ovoviviparous species appearas sister-taxa. This is confirmed by a molecular phylogeneticanalysis of the same species, based on the sequence of a portionof the 16S ribosomal RNA mitochondrial gene. Neither analysisunequivocally confirms the monophyly of Tectarius. The divergenceof DNA sequences within Tectarius suggests that the genus arosein the Upper Cretaceous, much earlier than the oldest (UpperEocene) fossils. Only 4 of the 175 species of Littorinidae areknown to be ovoviviparous (with brooding through metamorphosis)and the possible adaptive significance of this type of developmentis discussed. Hitherto, its rarity had been explained by earlyextinction of poorly-dispersed brooding taxa. However, ovoviviparitymay have persisted in Echininiopsis for at least 35 millionyears, and has not precluded colonization of islands 6300kmapart. (Received 17 September 1996; accepted 15 November 1996)     

MORPHOLOGICAL STUDIES OF LYMNAEID SNAILS FROM THE HUMAN FASCIOLIASIS ENDEMIC ZONE OF BOLIVIA

The taxonomic position of the Lymnaeidae from the BolivianAltiplano has been unclear. On the basis of conchological characters,some authors reported two species from this area, Lymnaea viatrix andL. cubensis while others, considering also anatomical characters,considered L. viatrix as a synonym of L. cubensis. More recentstudies demonstrated genetic identity between the Bolivian lymnaeidsand L. truncatula from the Iberian Peninsula. Populations recognizedas L. cubensis correspond to a distinct genetic group, but geneticinformation was not available for L. viatrix. In the light ofthese genetic results, a morphometric study of both the shell (usingRaupian parameters) and male reproductive system was carried outof L. cubensis from Cuba (type locality), Dominican Republic,Guadeloupe and Venezuela, and of L. truncatula from Bolivia,France, Portugal, Spain and Morocco. Syntypes of L. viatrixfrom Argentina (var. A. ventricosa) and specimens of L. viatrix(var. B. elongata) from the type locality, Peru, were also studied(conchological characters only). The conchological study showedthe presence of a large amount of variability between populations.This variability was not congruent with genetic results. Alarge amount of variability was also found using anatomicalcharacters of the male reproductive system and all of them clearlyseparate L. cubensis from L. truncatula independently from geographicalorigin. Thus anatomical characters, unlike conchological parameters,do differentiate taxonomic species inferred from genetic studies. (Received 11 September 1997; accepted 5 March 1999)     

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;     

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 GENETIC ANALYSIS OF SYMPATRIC SUBPOPULATIONS OF THE SIBLING SPECIES LITTORINA SAXATILIS (OLIVI) AND LITTORINA ARCANA HANNAFORD ELLIS

A detailed analysis of shell characteristics (four parameters)and allozyme allele frequencies (eight to nine polymorphic loci)of sympatric Littorina saxatilis and Littorina arcana subpopulationsfrom Filey Brigg on the east coast of England supports theiridentification as closely related but separate species. Oneenzyme system, esterase-2, was nearly species diagnostic. Resultswere generally comparable with an earlier study of the two speciesfrom East Lothian, Scotland. Littorina saxatilis was again foundto be more variable than L. arcana. (Received 14 December 1983;     

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;     

GENETIC AND CONCHOLOGICAL COMPARISON OF SNAILS (HELIX ASPERSA) DIFFERING IN SHELL DEPOSITION OF LEAD

Populations of snails inhabiting areas with different historiesof Pb contamination differed in their deposition of Pb in shellrelative to soft tissues. Genetic variation, measured usingisozymes, was not related to Pb history nor geographic distancebetween populations. Shell characteristics were significantlydifferent among sites; shell dry weight was strongly relatedto soil calcium levels. Shells of snails from areas with longhistories of Pb contamination were significantly more robust(greater shell width/shell height ratio) than snails from otherlocations. H. asprsa adaptation to Pb contamination may involvesignificant changes in shell characteristics but these do notcorrelate with genetic traits assessed with allozymes (Received 29 December 1994; accepted 15 October 1995)     

COMPARISONS OF THE BIOLOGY OF THE INTERTIDAL SUBANTARCTIC LIMPETS NACELLA CONCINNA AND KERGUELENELLA LATERALIS

Two limpet species occur intertidally on subantarctic SouthGeorgia, the patellid Nacella concinna and the siphonarlid Kerguelenellalateralis. N. concinna is confined to the lower shore closeto LWS; K. lateralis occurs in middle shore pools, so theirdistributions do not overlap. N. concinna has a much narrowerthermal niche (–12.9°C to +15.6°C) than K. lateralis(–17.8°C to +31.8°C). Environmental data are presentedto show that the upper lethal temperature of N. concinna islow enough to prevent the limpet living higher on the shore.Both limpet species are slow-moving, but K. lateralis showsincreasing speed with rising temperature, peaking at 15–20°C.In contrast, N. concinna moves actively down to –1.9°C(when sea water freezes), but there is a steady decrease inspeed of locomotion above +2°C. Locomotion ceases at 14°Cin N. concinna (c.f. 30°C in K. lateralis). Both speciesexhibit very low tenacities, but in N. concinna tenacity decreaseswith increasing shell length. In K. lateralis there is no effectof temperature on tenacity. Both species show a positive allometricrelationship between foot area and shell length. N. concinnafeeds upon microbial films and microepiflora, but K. lateraliseats colonial diatoms and Enteromorpha bulbosa. Observationson shell middens of the kelp gull Larus dominicanus showed thatthe gulls did not eat K. lateralis, though they ate great quantitiesof the less accessible N. concinna. Gulls ate N. concinna assmall as 11 mm shell length (within the size range of K. lateralis).Experiments on gulls demonstrated an unwillingness to eat K.lateralis, probably because the siphonariid extrudes a viscidwhite mucus when the foot is touched. (Received 9 May 1996; accepted 8 July 1996)     

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)     

FUNCTIONAL MORPHOLOGY AND DESCRIPTION OF A NEW SPECIES OF AMYGDALUM (MYTILOIDEA) FROM THE OXYGEN MINIMUM ZONE OF THE ARABIAN SEA

A new species of Amygdalum (Amygdalum anoxicolum n. sp.) isdescribed living in the soft green muds of the oxygen minimumzone off the Oman margin in the northern Arabian Sea. It isdistinguished by both its shell and anatomical characters. Theanatomy is described and discussed in relation to the environment.The presence of haemoglobin is viewed as a direct adaptationto the low levels of oxygen found in its habitat. The observedwide size range of ingested food particles is discussed in relationto the low oxygen environment. The nest building habit is assumedto be a key adaptation to living in soft, high porosity mud.The distribution of the genus is examined in relation to thedistribution of known zones of hypoxia in the world9s oceans,but no relationship is evident. Amygdalum species are not indicatorsof low oxygen environments. (Received 17 August 2000; accepted 14 November 2000)