SURFACING AND WATER LEAVING BEHAVIOUR OF THE FRESHWATER PULMONATE SNAILS LYMNAEA PEREGRA (MULLER), BIOMPHALARIA GLABRATA (SAY) AND BULINUS JOUSSEAUMEI (DAUTZENBERG)

Surfacing and water-leaving behaviour (quitting) of aquaticpulmonate snails was investigated in the laboratory. Regressioncoefficients of surfacing rate per hour for 16 snails in 200cm³ of water were 1.28 for L. peregra, 1.35 for B. glabrata(albino form), 0.47 for B. glabrata (pigmented form) and 0.17for B. jousseaumei. Disturbance, snail size and conditioningof the water were relatively unimportant factors; water volumeadjusted by depth was also unimportant up to 50 mm depth. Importantfactors controlling surfacing appeared to be population density,time of year, temperature, water volume adjusted by perimeterand dissolved oxygen concentration. The response to the latterwas not clear and both B. glabrata and L. peregra orientatedalmost as well to a nitrogen/water interface as to an air/waterinterface. Water-quitting regressed on volume, which had beenadjusted by perimeter variation, revealed a curvilinear relationshipfor B. glabrata, with maximum quitting at 1 snail per 7cm³ (1snail per 14cm of climbable perimeter). L. peregra showed anegative exponential curve, with maximum quitting at 1 snailper 3 cm³ of water (1 snail per 8 cm of climbable perimeter).These data could be relevant to design and management of irrigationchannels where bilharzia might occur. (Received 28 November 1990; accepted 22 October 1991)     

ALLOZYME VARIATION IN SOME POPULATIONS OF THE FRESHWATER SNAILS LYMNAEA PEREGRA, L. AURICULARIA AND L. STAGNALIS (GASTROPODA: PULMONATA)

Three species of the genus Lymnaea (Gastropoda, Pulmonata, Basommatophora)were examined for genetic variation at 11 enzymatic loci insamples collected in Brittany (North-Western France). Variabilitywas greatest in L. percgra, but the limited area of samplingdid not allow specific inferences. The UPGMA clustering of unbiasedgenetic distances (Nei, 1978) isolated first L. auriculariasamples, and secondly the single L. stagnalis sample from theL. peregra set. This scheme of relationships was opposed tothe identification of alleles shared by the three species atthe studied loci Analysis of L. peregra population structure by F-statistics(Weir & Cockerham, 1984) suggested high inbreeding and raisedthe possible role of self-fertilization. The contradiction betweenhigh variability and high homozygosity is discussed. Furthermore,population differentiation was rather weak (although statisticallysignificant), despite some remarkable differences among loci.Nm estimates using Wright's (1951) and Slatkin's (1985) methodsrevealed a gene flow unable to prevent the effect of geneticdrift among populations. The relative homogeneity of populationswith weak genetic exchange is discussed (Received 24 May 1993; accepted 21 March 1994)     

POPULATION GENETICS OF LYMNAEA PEREGRA (MULLER) (GASTROPODA: PULMONATA) IN LAKE GENEVA

Using gel electrophoresis, a study was made of genetic exchangeamongst populations of Lymnaea peregra in Lake Geneva. Geneticvariability is high. A within-population heterozygote deficiencyis shown and the role of null alleles as an explanation of thisdeficiency is discussed. High values of gene flow show thatthe inter-population structure is weak. Exchange between populationspropably occurs by both active and passive means. (Received 9 January 1989; accepted 15 August 1989)     

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;     

BARNACLE-DWELLING EGOTYPES OF THREE BRITISH LITTORINA SPECIES AND THE STATUS OF LITTORINA NEGLECTA BEAN

On the basis of collections from SW Wales and W Ireland it isshown that small (<5 mm) black and/or white Littorina occurringin the barnacle zone on exposed shores, of the L. neglecta shelltype, comprise three species. From their reproductive anatomythese correspond to L. saxatilis, L. arcana and L. nigrolineata,and are interpreted as analogous barnacle-dwelling ecotypesof these species. This adds to the evidence that L. neglectais not a distinct species, but an ecotype of L. saxatilis. (Received 1 June 1992; accepted 23 June 1992)     

INTERPOPULATION VARIATION IN JUVENILE SURVIVAL AND GROWTH RATES OF LYMNAEA PEREGRA (GASTROPODA: PULMONATA); TEMPERATURE AT RECRUITMENT AS A SELECTION PRESSURE?

Three neighbouring populations of Lymnaea peregra had recruitmentin the summer (June and July), but one population (Sheaf) hada second recruitment in September and October. We hypothesizedthat juveniles of the Sheaf population would be subject to selectionunder both ‘summer’ and ‘winter’ conditions,and thus should be more resistant to low-temperature stressthan juveniles of the other populations. The hypothesis wassupported by the findings that Sheaf juveniles survived andgrew better over a wider range of temperatures (2, 10, 15 and20°C )while juveniles of the other two populations wereadapted only to higher temperatures (15 and 20°C). Therewas evidence that some of these traits were genetically fixed. ^*Present address: Department of Applied Science, City Polytechnicof Hong Kong, Mongkok, Kowloon, Hong Kong. (Received 15 September 1988; accepted 16 December 1988)     

ALLOZYME SIMILARITY OF ATLANTIC AND PACIFIC SPECIES OF LITTORINA (GASTROPODA: LITTORINIDAE)

Four Littorina species from the Pacific (L. sitkana, L. brevicula,L. mandshurica, L. squalida) and four from the Atlantic (L.mariae, L. obtusata, L. saxatilis and L. littorea) were comparedelectrophoretically using 17 loci. Analysis of genetic identities showed that there were threegroups. The first group comprises L. sitkana, L. mariae, L.obtusata and L. saxatilis The second group consists of L. breviculaand L. mandshurica and the third one includes L. squalida andL. littorea. Our results support Reid's hypothesis of the originof Atlantic littorines from two Pacific ancestors. (Received 18 December 1991; accepted 30 March 1992)     

THE SYSTEMATIC POSITION OF LAROCHEA FINLAY, 1927, AND INTRODUCTION OF A NEW GENUS AND TWO NEW SPECIES (GASTROPODA: SCISSURELLIDAE)

Species of the gastropod genus Larochea Finlay, 1927 are shownto be scissurellids without an anal shell slit or foramen. TheNew Zealand species, L. miranda Finlay, 1927 and L. secundaPowell, 1937, brood their young in the right subpallial cavityagainst a modified inner lip that is set well behind the aperturalplane. Larochea scitula n.sp. is based on shells from WanganellaBank, southern Norfolk Ridge. Larocheopsis n. gen. is introducedfor a minute species from off northern New Zealand that lacksa shell brood chamber. Larochea miranda and Larocheopsis amplexan.sp. are either gonochoristic with smaller males or consecutivehermaphrodites, while Larochea secunda and L. scitula are evidentlygonochoristic, having mature males and females of similar size. (Received 23 July 1992; accepted 10 December 1992)     

EMBRYONIC ABNORMALITIES IN POPULATIONS OF LITTORINA SAXATILIS (OLIVI) (GASTROPODA: PROSOBRANCHIA) IN THE WHITE SEA

The incidence of abnormally developing embryos in brood pouchesof females was studied in five populations of Littorina saxatilisliving under different salinity conditions. The proportion ofabnormals decreased during embryonic development. Age of femaleand infection with the ciliate Protophrya ovi-cola had no effecton the frequency of abnormals. The proportion of abnormal eggs(PAE) increased at the end of the reproductive season; a similartrend was found for the proportion of abnormal veligers (PAV)although it was not always significant. The proportion of abnormaljuveniles (PAJ) showed no seasonality. The increase of PAE coincidedwith decrease of water temperature in September. Environmentalsalinity was significantly correlated with frequency of abnormalsin L. saxatilis populations: the highest values of PAV and PAJwere observed in estuarine population (salinity range 5–8%),intermediate in populations living under conditions of 10–20%osalinityand lowest in control populations (23–24%). Itis suggested that critical salinity (sensu Khle-bovich, 1974)(approximately 8–10%) may also be a crucial point forstability of embryonic development in the White Sea populationsof L. saxatilis. (Received 28 June 1994; accepted 28 November 1994)     

OBSERVATIONS ON THE ECOLOGY OF BULINUS (PHYSOPSIS) GLOBOSUS (MORELET), B. (BULINUS) FORSKALII (EHRENBERG) AND LYMNAEA NATALENSIS (KRAUSS) IN RICE SWAMPS IN SIERRA LEONE.

Changes in population of Bulinus globosus, B. forskalii andLymnaea natalensis were studied in four different locationsnamed Sites A, B, C. and D in Bo town Sierra Leone. The snailpopulations and oxygen concentrations of water increased inthe rainy season. The biological oxygen demand did not havesignificant correlation with changes in snail populations inSites B and D. B. globosus was absent from Site A while B. forskaliiwas the predominant snail in Site C. Site D with the highestmean monthly conductivity of 176 µmhos contained 54% whileSite A with a similar reading of 90µmhos contained 1%of all snails found. (Received 22 May 1980;     

THE FINE STRUCTURE OF DIFFERENTIATING SPERMATOZOA AND SERTOLI CELLS IN THE GONAD OF THE POND SNAIL, LYMNAEA STAGNALIS

The fine structure has been examined, of spermatogonia, spermatocytes,early spermatids, late spermatids and early spermatozoa nestlingagainst Sertoli cells in the gonad of Lymnaea stagnalis. Changes in the Sertoli cells are linked with the phases of spermdifferentiation. Details on differentiation particularly ofthe head of the sperm, are presented. (Received 14 March 1981;     

ULTRASTRUCTURAL AND ECOLOGICAL ASPECTS OF THE DEVELOPMENT OF CHLOROPLAST RETENTION IN THE SACOGLOSSAN GASTROPOD ELYSIA TIMIDA

Elysia timida is a common and endemic inhabitant of shallowand very well lit waters in the Mediterranean. This sacoglossanslug retains functional symbiotic chloroplasts derived fromits algal food, Acetabularia acetabulum, although the chloroplastsare not transmitted in the spawn. After hatching and until day12, Elysia timida juveniles do not retain these chloroplastsin the digestive gland. However, newly hatched juveniles retainchloroplasts from Cladophora dalmatica. Development varies seasonallybetween direct (December to April) and lecithotrophic (October,November and May), and this variation may be an adaptation toseasonal calcification of the algal food Acetabularia acetabulum. (Received 12 February 1991; accepted 15 July 1992)     

ECOLOGY OF THE INTRODUCED SNAIL MELANOIDES TUBERCULATA (GASTROPODA: THIARIDAE) IN RELATION TO BIOMPHALARIA GLABRATA IN THE MARSHY FOREST ZONE OF GUADELOUPE, FRENCH WEST INDIES

Melanoides tuberculata was introduced to several types of aquatichabitat occurring in the marshy forest zone, Guadeloupe, asa competitor of the snail host, Biomphalaria glabrata, of Schistosomamansoni. Colonization by the competitor was successful in allhabitats, but the impact on the snail host was variable accordingto the site. Evolution of global prevalences, intensities andabundances of Schistosoma mansoni in the reservoir host, Rattusrattus, did not show any significant changes during the eightyears following the introduction of the competitor. The analysisof size distribution of M. tuberculata in a release plot showedthat maximum reproduction took place during the rainy seasonbetween June and November. The thiarid snail needed 4.5 monthsto reach a size of 10 mm, 14 months to reach 15 mm, and morethan 30 months to reach 20 mm. Calculated life expectancieswere 0.74 year at the first year of age, 0.9 year at the secondyear, and 0.5 year at the third year. (Received 21 September 1992; accepted 5 January 1993)     

THE OLFACTORY-GUSTATORY BASIS OF FOOD PREFERENCE IN THE HERBIVOROUS PROSOBRANCH, LITTORINA LITTOREA (LINNAEUS)

Studies of food preference often measure the rate of consumptionof prey items by a predator; this confounds the factors determiningattractiveness and edibility, ignoring the fact that preferencesare probably resolved at the pre-ingestive stage of a consumer'sfeeding behaviour. In this paper, a novel technique has beendeveloped to measure the importance of chemical stimuli to feed,perceived by olfactory and gustatory means, as determinantsof the subsequent feeding behaviour of the generalist intertidalherbivore,Littorina littorea (L.). Cell-free extracts of a rangeof algae were presented simultaneously, impregnated in artificialcellulose-based substrates. Subsequent ranking of feeding preferencesreveals hierarchies which correspond with those establishedby other workers, using intact algae to feed L. littorea. Extractsof ephemeral early successional species (e.g. Ulva lactuca,Porphyra umbilicalis) generally provoke a more positive feedingresponse than perennial late successional species (e.g. Fucusvesiculosus, Asco-phyllum nodosum). Animals fed a preferredalga (Ulva lactuca) prior to preference trials displayed a reductionin the strength of preference relative to starved animals. Therewas no evidence of ‘switching’ of preference rankingsdue to satiation, however. These findings confirm the importanceof olfactory-and gustatory-mediated feeding behaviour as theprimary determinant of food preference in L. littorea. Theyare consistent with the hypothesis that the factors which determinethe attractiveness of a food type, such as chemical cues, arethe criteria by which consumers predict the value of that food.     

FOOD-RELATED CONDITIONING AND NEURONAL CORRELATES IN THE FRESHWATER SNAIL LYMNAEA STAGNALIS

The feeding response of Lymnaea stagnalis shows robust appetitiveconditioning to a novel chemostim-ulus (amyl acetate), whichis retained for more than 4 days. In scrru-isolated centralnervous system (CNS)-hp preparations taken from conditionedsnails, application of amyl acetate to lip tissue led to theonset of fictive feeding in 5/17 individuals, and excitationof an identified cerebral-buccal feeding intemeuron, CV1_a. ina further 8. Control group snails showed no response to amylacetate. Attempts were made to aversively condition L. stagnalis usinga food stimulus (sucrose) as the conditioned stimulus and eitheran electric shock or mechanical stimulus as the unconditionedstimulus. No conditioned response to sucrose (i.e. withdrawal)was seen following training. However, trained snails exhibiteda reduction in responsiveness which was manifest as inhibitionof the feeding response to sucrose and an increase in the timetaken to emerge from the shell following handling. This reducedresponsiveness was seen at 1 hr but was lost by 24 hr aftertraining, and was not observed in control group snails. Semi-isolatedCNS-hp preparations from experimental snails recorded 1-3 hrafter training exhibited either inhibition of fictive feedingor no response following application of sucrose. Control groupsnails showed the normal excitatory response to sucrose includinginduction of feeding motor output Differences in food-related learning between Lymnaea and othergastropod species are discussed in relation to lifestyle andfeeding strategies (Received 14 January 1996; accepted 3 May 1996)     

RECRUITMENT AND GROWTH OF THE BLUE-RAYED LIMPET, HELCION PELLUCIDUM (L.), IN SOUTH EAST IRELAND

This account reports on recruitment, growth and population structureof the macroherbivorous gastropod Helcion pellucidum (L.) onan exposed rocky shore in S.E. Ireland, with particular referenceto the pellucidum form. Recently settled Helcion spat were foundinitially on living crustose lithothamnia in pools and on openrock, on the lower shore, during February-April and subsequentlyrecruited to macro-algal fronds. By May they were found in abundanceon Mastocarpus stellatus and Laminaria spp. Helcion was markedlyseasonal in its occurrence on Mastocarpus, being most abundantduring May-August. A complementary pattern was evident on growingHimanthalia receptacles, with limpets abundant from July onwards.It is suggested that Helcion may migrate from Mastocarpus toHimanthalia as they grow. Data on growth rates are presentedfor Helcion in the various habitats studied. Growth rates werehighest on the Laminaria spp. (Received 21 February 1992; accepted 8 June 1992)     

THE FINE STRUCTURE AND FUNCTION OF THE ANAL GLAND OF THE MURICID NUCELLA LAPILLUS (NEOGASTROPODA) (AND A COMPARISON WITH THAT OF THE TROCHID GIBBULA CINERARIA)

The fine structure of the anal gland of Nucella lapillus isdescribed, and compared with that of Gibbula cineraria. It isconcluded that the glands are not analogous in function andare unlikely to be homologous. The anal gland of Gibbula isa small rectal diverticu-lum near the anal papilla, lined bymucus-secreting cells. The evidence suggests that the largeanal gland of Nucella extracts macromolecules and cations fromthe blood, metabolizes or sequesters them in lyso-somes andultimately expels the resultant residual bodies by apocrinesecretion. It has large reserves of lipid and glycogen, andthe abundance of melanin indicates that it is a major site oftyrosine degradation. The invariable presence of bacteria inthe lumen, and of pits in the epithelium to house them, impliesa symbiotic relationship in which the bacteria metabolize anddegrade the cell debris, some of which is resorbed. The lossof the anal gland in bucci-nids and nassariids is accompaniedby great reduction in size and importance of the rectal sinus,and increase in complexity of the kidney. (Received 31 December 1991; accepted 23 January 1992)     

THE GENETIC RELATIONSHIPS OF THREE TAXA IN THE LITTORINA SAXATILIS SPECIES COMPLEX (PROSOBRANCHIA: LITTORINIDAE)

Three species in the Littorina saxatilis complex (the ovoviviparousL. saxatilis and the two oviparous species L. arcana and L.nigrolineata) were screened for the products of up to 22 lociusing gel elec-trophoresis. Analysis of allele frequencies andderived genetic distances showed that, with rare exceptionsin L. saxatilis and L. arcana, conspecific populations clusteredtogether. Sympatric pairs of populations showed significantdifferentiation at many polymorphic loci. The results confirmedthe biological reality of the three taxa, and did not suggestthe existence of any previously unrecognized species. The speciesare genetically closely related to each other, with Nei geneticdistances ranging from 0.035 to 0.083. (Received 24 October 1989; accepted 18 April 1990)     

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;