INVESTIGATIONS OF THE ROLE OF MUCUS AND FAECES IN INTERSPECIFIC INTERACTIONS OF TWO LAND SNAILS

A previous report (Smallridge & Kirby, 1988) showed interspecificcompetition between the two land snails Theba pisana and Cernuellavirgata and suggested the mechanism of competition may be thatheterospecific mucus and excrement inhibits activity. We describea series of experiments in which we could find no consistentevidence for any short-term inhibition of activity in the presenceof heterospecific mucus. Furthermore snails did not consistentlyavoid mucus-treated surfaces or mucus-coated food. Cernuellastopped at and fed from snail faeces (both conspecific and heterospecific)more than did Theba. This suggests an alternate mechanism forinterspecific competition. (Received 9 April 1992; accepted 9 June 1992)     

EVOLUTION, DURING DIGESTION, OF THE BACTERIAL FLORA IN THE ALIMENTARY SYSTEM OF HELIX ASPERSA (GASTROPODA: PULMONATA): A SCANNING ELECTRON MICROSCOPE STUDY

The use of scanning electron microscopy (SEM) allowed a studyof the distribution of bacteria in the various digestive organsof the snail Helix aspersa Müller. The bacteria are enclosedby mucous secretions (mucous film or mucous grains) and sometimesattached on the cilia of some of the digestive walls. Accordingto the food that was given to the snails, different morphologicaltypes appeared, two of which dominated. Adult snails were fasted for 4 days, given a dehydrated artificialfood and then sacrificed at different times during digestion.The presence of bacteria may be related to the time of digestion.In fact, bacteria seem to accompany the food mass; they developmostly in the stomach and in the intestine where they may helpdigest the food. Fasting or hibernating snails do not possess bacteria in thealimentary lumen or on the digestive walls. However, the residualfaeces localized in the distal, intestinal lumen, lodge greatquantities of bacteria. From these results, the endogenous or/and exogenous existenceof the bacterial flora in alimentary system of Helix aspersais discussed. (Received 26 June 1989; accepted 16 October 1989)     

POPULATION DENSITY EFFECTS ON GROWTH IN CULTURE OF THE EDIBLE SNAIL HELIX ASPERSA VAR. MAXIMA

The effects of population density on the growth of H. aspersaMÜller var. maxima under controlled environmental conditionswere examined. Inhibitory effects on snail growth and maturityresulting from increased population density, between 100–800snails m^–2 of floor area, were observed for a range ofcontainer cleaning frequencies. At all population densities,enhanced snail growth was observed when the frequency of containercleaning was increased to a two-day interval. No significantdifferences were recorded, following 19 weeks growth, betweenfinal mean weights of snails from containers cleaned less frequently.The lowest snail mortality was consistently recorded at thelowest population density in the most frequently cleaned containers.At all snail population densities three phases of growth wereobserved: (a) lag (0–5 weeks), (b) rapid (6–15 weeks)and (c) stable (16 weeks and over). During the first three weeksof growth, high population density had a positive effect ondiet consumption, food conversion efficiency and snail growth.Adverse population density effects increased progressively duringphase (b), typically following 9 weeks growth. Juvenile snailstransferred from high to low population densities during phase(b) continued to exhibit slower growth rates associated withhigh population densities. Food conversion efficiency of snailsin all treatments decreased throughout the experimental periodbut with no overall effect of container cleaning frequency apparent.Inherent growth variability of sibling snails was unaffectedby population density or container cleaning frequency. The importanceof the results for intensive snail culture is discussed. (Received 23 June 1994; accepted 1 December 1994)     

GROWTH AND DEMOGRAPHY OF POPULATIONS OF BIOMPHALARIA PFEIFFERI (GASTROPODA, PLANORBIDAE) IN THE LABORATORY

The growth and demography of the freshwater snail Biomphalariapfeifferi, an intermediate host of the parasitic worm Schistosomamansoni in Eastern Zaire, were investigated in the laboratoryunder two conditions of crowding. Both individual and populationgrowth were depressed by crowding. Crowding reduced growth rateand all the variables and parameters related to reproduction(fecundity, instantaneous birth rate, net reproduction rate),but did not affect maximum size, survivorship, instantaneousdeath rate and life expectancy. The mechanism involved in theseeffects is likely to be exploitation competition for especiallyprofitable food items such as diatoms. (Received 10 November 1986;     

THE INFLUENCE OF INGESTIVE CONDITIONING ON FOOD CHOICES IN THE LAND SNAIL HELIX ASPERSAMULLER (GASTROPODA: PULMONATA: STYLOMMATOPHORA)

An experiment of food choice in one-day-old naïve landsnail Helix aspersa Müller was carried out, involving threephases. First, a training period on a monophagous diet of leafdiscs of Taraxacum officinale or Urtica dioica was given for1, 5, 10, 15 or 30 days. The area of leaf discs consumed byeach snail during this stage was assessed. In the intermediatephase, the animals were not fed for 24hours. Finally, food choicewas tested over 24 hours, when animals could eat leaf discsof each species of plant usedfor training. A coefficient offood preference for Urtica dioica was calculated (UPC = areaof Vrtica leaf disc consumed/area of Urtica and Taraxacum discsof leaves consumed for 24 hours). The length of the training phase did not reduce the percentageof individuals preferring the plant previously consumed. Onthe contrary, after 15 days of monophagous diet, animals formerlyfed on Urtica showed a coefficient of food preference for Urticasignificantly higher than Taraxacum fed animals, and after 30days of training this phenomenon was more pronounced. Thus,during the choice test, snails ate more of the previously eatenplant leaf. Our experiment highlighted the feeding behaviourplasticity of H. aspersa. The significant correlation betweenthe area of Urtica eaten during the training phase and the coefficientof food preference for Urtica is discussed. (Received 10 April 1994; accepted 10 January 1995)     

BIOCHEMICAL COMPOSITION OF HELIX SNAILS: INFLUENCE OF GENETIC AND PHYSIOLOGICAL FACTORS

This paper describes the biochemical composition of differentspecies (Helix lucorum, Helix pomatia) and sub-species of snails(Helix aspersa aspersa, Helix aspersa maxima) reared in thesame conditions with a feed (‘Helixal’) speciallydesigned for edible snails. In addition, the composition ofwild H. pomatia and H. lucorum is presented to allow comparisonbetween snails of different origins. Analyses determined thepercentages of proteins, lipids and minerals. They reveal bothsimilarities and differences in composition according to thespecies and the part analysed (whole body, pedal mass, and visceralmass). H. pomatia contains the highest percentage of mineralmatter and the lowest percentage of lipids. Surprisingly, proteincontents are slightly different between artificially rearedH. aspersa maxima of 3 months old and wild H. pomatia. The resultsmake it possible to evaluate nutritional quality of snails withthe composition of the body of four edible snail species. (Received 16 September 1996; accepted 24 May 1997)     

THE RELATIONSHIP OF BODY SIZE AND GROWTH TO EGG PRODUCTION IN THE HERMAPHRODITIC FRESHWATER SNAIL, HELISOMA TRIVOLVIS

Among several other factors, body size has been found to influenceegg production in several species of hermaphroditic snail. Wetested whether this relationship between body size and egg productionexists in Helisoma trivolvis, a freshwater hermaphroditic species.We isolated 50 H. trivolvis from a laboratory population, measuredshell diameter, and monitored egg production for seven weeks.We found a positive relationship between body size and totalnumber of eggs produced, as well as body size and number ofeggs per egg mass. When body size and egg production are linked,it should be adaptive for larger individuals to act as femalesand smaller individuals as males. Since body size is relatedto female fecundity in this species, the relative size of snailsshould determine, at least in part, which individual acts asmale and which as female during copulation. However, the relationshipbetween body size and egg production is not nearly as strongas it is in other snail genera. Other factors such as age, genotypeand previous experience may be important in determining egg-layingcapacity and therefore gender choice in this species. In addition,we found a negative relationship between growth during thisperiod and egg production. This relationship has been foundin other pulmonates, and is evidence of resource allocationtradeoffs. (Received 3 August 2004; accepted 2 August 2005)     

FIELD OBSERVATIONS ON FEEDING OF THE LAND SNAIL HELIX ASPERSA MULLER

Feeding of the land snail Helix aspersa (Müller) was observedat monthly intervals. Three natural populations in Galicia (NW-Spain)were studied. At two sites only a few plants constituted thebulk of the diet and in spring the snails' diet had the highestdiversity (H'). In the third population feeding and distributionof Helix aspersa (Müller) were observed in a small plotwith permanent patches of Urtica dioica. Nearly one half offeeding snails fed upon Urtica dioica. Most of the other observations wereon Mentha suaveolens, Ranunculus repens and Gramineae. The diversity ofthe snails' diet showed seasonal variation with the maximumin the autumn months. Comparison between the availability ofthe different plant species and their contribution to the snails'diet showed that the snails did not eat at random; Urtica dioicawas eaten much more than expected from its occurrence and grasseswere strongly under-represented in the snails' diet. Temporalchanges of availability were significantly correlated with the amountseaten in the case of Urtica, but not for the other food plants.The distribution of the snails in the plot was significantlycorrelated with that of Urtica. Chemical analyses of the foodplants revealed Urtica as the species with the higher protein,ash and calcium contents. The strong preference of Helix aspersafor Urtica dioica could be explained by the value of Urticaas food or by its suitability as habitat for the snails. The largestproportions of green material in the snails' diet occurred inthe spring and juveniles ate more green material than adultsin the three populations. (Received 16 March 1998; accepted 30 November 1998)     

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)     

OBSERVATIONS ON THE POPULATION DYNAMICS OF SNAIL HOSTS FOR SCHISTOSOMES IN THE DELTA OF THE SENEGAL RIVER BASIN

Between 1987 and 1990, snail surveys were carried out in theSenegal River Basin at Lampsar, in a natural ‘marigot’,and at Richard Toll, in an irrigation canal. Lymnaea natalensis.Bulinus guernet^* and B.forskalii, were regularly found at boththe sites. Bulinus globosus/jousseaumei and B. umbilicatus wereonly present at Lampsar. A decrease of the snail density wasobserved during the rainy season at Lampsar. The recent appearanceof Biomphalaria pfeifferi at Richard Toll is discussed. Thedata show that adult B. pfeiffert are present in the main irrigationcanal all through the year. Consequently parasite transmissionwill be possible throughout the year as well. (Received 23 October 1991; accepted 20 May 1992)     

OTHER PAPERS: INTERFERENCE AND RESOURCE COMPETITION IN TWO LAND SNAILS: ADULTS INHIBIT CONSPECIFIC JUVENILE GROWTH IN FIELD AND LABORATORY

Interference and resource competition by adults inhibited growthrates of conspecific juveniles of the land snail species Mesodonthyroidus and Neohelix albolabris in separate field and laboratoryexperiments, but not in laboratory experiments on Anguispiraalternata. In 1 m² field cages at near-natural densities underambient food and water conditions, juvenile M. thyroidus apparentlycompeted with adults for food or water or both resources, growingmore slowly when living with two conspecific adults, but beingunaffected by adult presence when food and water were augmented.Neohelix albolabris juveniles were similarly unaffected in fieldcages by presence of two adults when food and water were augmented.In contrast, interference, not resource competition, apparentlyexplained growth inhibition in laboratory cages at densitiesconsiderably greater than natural densities, with non-limitingfood and moisture; both M. thyroidus and N. albolabris juvenilesgrew more slowly as conspecific adult number increased fromzero to three. (Received 17 July 1995; accepted 11 November 1996)     

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)     

ANATOMY AND SYSTEMATIC PLACEMENT OF SAYELLA LAEVIGATA (GASTROPODA: PYRAMIDELLIDAE: SAYELLINAE)

The ectoparasitic snail Sayella laevigata was first describedby Orbigny (1842) and first placed in the genus Sayella by Odé (1994) without discussion. This study provides justificationfor this assignment to the genus Sayella Dall, 1885 and to the subfamilySayellinae Wise, 1996. Sayella laevigata possesses an elongatepenis, with subapical swelling tapering to a short terminalnipple, that is characteristic of the genus.     

INFLUENCE OF THE QUINOLIZIDINE ALKALOID CONTENT OF LUPINUS ALBUS (FABACEAE) ON THE FEEDING CHOICE OF HELIX ASPERSA (GASTROPODA: PULMONATA)

Adult Helix aspersa snails were maintained individually forone week in plastic cages with 9 living Lupinus albus plantsas their only food. Among these 9 plants, 3 chemotypes bitter,intermediate and sweet which differed in their alkaloid contentwere equally represented. Each day, the leaf surface grazed and the number of leaves attackedby the snails were recorded for each chemotype and each snail.A consumption/ attack (C/A) ratio was calculated by dividing thesurface grazed (C) by the number of attacks (A). The numberof attacks and the grazed area were positively correlated foreach chemotype during the whole experiment, and the snails atesimilar quantities of lupin each day. After 4 and 6 days ofexperiment, we noticed a rejection of the bitter chemotype infavor of the intermediate and sweet ones respectively. Afterthe 6th day, the surface grazed per attack was significantlyhigher on the sweet chemotype than on the bitter plants. Wehypothesize that rejection of the bitter chemotype might berelated to (i) an alkaloids reaction threshold associated withan increase in the amount of alkaloids in the wounded plantsand/or (ii) aversive ingestive conditioning. (Received ; accepted 16 April 1999)     

A DEMOGRAPHIC ANALYSIS OF y-NEOTRICULA APERTA (GASTROPODA: POMATIOPSIDAE) POPULATIONS IN THAILAND AND SOUTHERN LAOS, IN RELATION TO THE TRANSMISSION OF SCHISTOSOMIASIS

The results of field work undertaken along the tropical reachesof the Mekong river in Thailand and Laos are presented. Demographicdata concerning the schistosome-transmitting snail y-Neotriculaaperta are examined and related to the life-cycle of this species.On the basis of these data, and a review of the literature,the only available, empirically substantiated, model for thelife-cycle of y-N. aperta (see Upatham et al., 1980) is shownto be equivocal. In this model snails survive the annual spatein the Mekong river as eggs, which remain dormant below theflood waters, whilst the adults die soon after the waters beginto rise. Here, the above described model of ‘egg survival’is shown to fail to balance the demographic equation for y-N.aperta. An alternative model of delayed oviposition with post-spatesurvival of mature snails is proposed and shown to be a betterfit to the available data. The implications of this alternativemodel for the evolutionary biology and ecology of N. apertaare discussed. The possible effects of the Pak-Mul dam project(Northeast Thailand) on local populations of schistosome-transmittingsnails are assessed and the hydrodynamic conditions conduciveto vector snail proliferation are considered. Attenuation ofthe normal flood-cycle of the Mekong river anc alteration ofthe river bed topography will probably lead to increased numbersof N. aperta at Pak-Mul. These changes, together with the influxof casual labour from endemic areas in Laos, may result in anepidemic of human schistosomiasis in Northeast Thailand. (Received 18 April 1994; accepted 23 June 1994)     

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)     

THE COURTSHIP OF EUGLANDINA ROSEA FERUSSAC

Courtship of Euglandina rosea commences with trail followingculminating in the pursuing snail mounting the shell of thepursued from the rear. A vigorous head waving display then occursand is terminated after about 15 min by a short quiescent periodduring which the lower animal turns its head to face its ownshell. Copulation takes place whilst one snail is still mountedon the shell of the other, but in a head to head position broughtabout by the twisting of the neck of the lower animal. It lastsup to four hours. (Received 24 April 1984;     

Prey size, prey nutrition, and food handling by shrews of different body sizes

We tested some predictions relating metabolic constraints offoraging behavior and prey selection by comparing food handlingand utilization in four sympatric shrew species: Sorex minutus(mean body mass = 3.0 g), S. araneus (8.0 g), Neomys anomalus(10.0 g), and N. fodiens (14.4 g). Live fly larvae, mealwormlarvae, and aquatic arthropods were offered to shrews as smallprey (body mass <0.1 g). Live earthworms, snails, and smallfish were offered as large prey (>0.3 g). The larvae werethe high-nutrition food (>8 kJ/g), and the other prey werethe low-nutrition food (<4 kJ/g). The smallest shrew, S.minutus, utilized (ate + hoarded) <30% of offered food,and the other species utilized >48% of food. The largerthe shrew, the more prey it ate per capita. However, highlyenergetic insect larvae composed 75% of food utilized by S.minutus and only >40% of the food utilized by the other species. Thus, inverse relationships appeared between shrewbody mass and mass-specific food mass utilization and betweenshrew body mass and mass-specific food energy utilization:the largest shrew, N. fodiens, utilized the least food massand the least energy quantity per 1 g of its body mass. Also,the proportion of food hoarded by shrews decreased with increase in size of shrew. With the exception of S. araneus, the sizeof prey hoarded by the shrews was significantly larger thanthe size of prey eaten. Tiny S. minutus hoarded and ate smallerprey items than the other shrews, and large N. fodiens hoardedlarger prey than the other shrews.     

EGG LAYING BEHAVIOUR OF BULINUS OCTOPLOIDUS BURCH (BASOMMATOPHORA: PLANORBIDAE)

Egg laying behaviour of Bulinus octoploidus in the laboratorycomprises three phases: (1) grazing, (2) deposition and (3)sweeping. The grazing phase lasts about 60–90 min andcan be recognized by the slowness of forward movement, movementin a restricted area, extended pseudobranch, and regular radularmovements against the substratum. Radular movements stop onlyduring turns and have a maximum frequency of about 45 min^–1.Egg mass deposition occurs at the end of the grazing phase andis accomplished in 1 min or less. Egg mass sweeping is a post-depositionphase lasting 8—12 min, during which the snail "sweeps"back and forth across the egg mass, apparently to anchor andflatten it. ¹ Authors' present address: Museum of Zoology, The Universityof Michigan, Ann Arbor, Michigan 48109, U.S.A. (Received 23 January 1979;