FIELD OBSERVATIONS ON FEEDING OF THE LAND SNAIL HELIX ASPERSA MULLER

Feeding of the land snail Helix aspersa (Müller) was observedat monthlyintervals. Three natural populations in Galicia (NW-Spain)were studied. At two sites only a fewplants 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 observationswereon Mentha suaveolens, Ranunculus repens and Gramineae. The diversityofthe snails' diet showed seasonal variation with the maximumin the autumn months.Comparison between the availability ofthe different plant species and their contribution to thesnails'diet showed that the snails did not eat at random; Urtica dioicawas eatenmuch more than expected from its occurrence and grasseswere strongly under-represented in thesnails' diet. Temporalchanges of availability were significantly correlated with theamountseaten in the case of Urtica, but not for the other food plants.The distribution ofthe snails in the plot was significantlycorrelated with that of Urtica. Chemical analysesof the foodplants revealed Urtica as the species with the higher protein,ash and calciumcontents. The strong preference of Helix aspersafor Urtica dioica could beexplained by the value of Urticaas food or by its suitability as habitat for the snails. Thelargestproportions of green material in the snails' diet occurred inthe spring and juvenilesate more green material than adultsin the three populations. (Received 16 March 1998; accepted 30 November 1998)     

GENETIC STUDIES OF ALLOZYME VARIATION IN LEUCINE AMINOPEPTIDASE IN THE LAND SNAIL HELIX ASPERSA (MULLER)

In the land snail H. aspersathe enzyme LAP has two loci, LAP-1and LAP-2, both of which arc monomeric enzymes under the controlof multiple alleles, the alleles being codominant. None of theobserved ratios in the pheno types in the experimental progenywere significantly different from Mendelian expectation. ^* Present Address: Bournside School, Cheltenham, Glos. (Received 1 September 1981;     

HETEROGENIC COPULATORY BEHAVIOUR PRODUCES NON-RANDOM MATING IN LABORATORY TRIALS IN THE LAND SNAIL HELIX ASPERSA MULLER

This study reports the results of mate-choice experiments betweenresident and non-resident individuals of the simultaneous hermaphonteland snail Helix aspersa. Snails from different sites differedin their mating proclivity, which resulted in non-random matingin laboratory mate-choice tests. Those snails with the highermating propensity therefore tended to mate with their own typeand tended to be chosen as partner because they were also activelyseeking mates. Keeping snails under crowded conditions did noteffect a lasting influence on mating behaviour in laboratorytests, despite the decline in copulatory behaviour under crowdedfield conditions. (Received 7 November 1994; accepted 31 August 1995)     

SNAIL FARMING: A SURVEY OF BREEDING MANAGEMENT, HYGIENE AND PARASITISM OF THE GARDEN SNAIL, HELIX ASPERSA MULLER

Fifteen farms in 1984 and twenty in 1985 were investigated forbreeding management, hygiene and parasitism. Farm hygiene didnot seem to play an important role on the breeding performancesin our samples. Nematodes were the most frequent parasites:Alloionema appendiculatum were equally prevalent among juvenileand adult snails whereas Angiostoma aspersa and Nemhelix bakeriwere found mostly in reproductive adults. The presence of nematodes,assessed by coproculture, was negatively related to breedingperformances     

VARIABILITY OF EGG CANNIBALISM IN THE LAND SNAIL HELIX ASPERSA IN RELATION TO THE NUMBER OF EGGS AVAILABLE AND THE PRESENCE OF SOIL

Egg cannibalism by hatchlings has been demonstrated in somepulmonate land snails; this behaviour is promoted by a highhatching asynchrony within the egg-batch. Under laboratory conditions,the percentage of new-born snails Helix aspersa having cannibalisedunhatched eggs was not influenced by the soil factor: about70% of them ingested one egg within their first four days oflife whether soil was present or not. The propensity to eggcannibalism in hatchlings of H. aspersa increased with egg density.However, most of the new-born hatchlings consumed a single eggduring the four days following hatching, and only exceptionallytwo. The consumption of one egg increased the snails wet weightby 38.7% within four days. A weak ingestion of soil componentsalso occurred, but it induced a growth that was three-timesless than that due to the consumption of an egg. In addition,the survival of newly hatched snails maintained under non-dehydratingthermohygrometric conditions was high, even when they were submittedto four days food-deprivation. (Received 22 July 1999; accepted 24 November 1999)     

THE LOVE DART OF HELIX ASPERSA MULLER IS NOT A GIFT OF CALCIUM

The phenomenon of dart shooting in several species of land snailshas still not been explained. We were interested in whetherthe dart can function as a nuptial gift of calcium, as previouslyproposed. Donating calcium would increase the fitness of theoffspring and thereby result in a higher reproductive successfor the donor. We confirmed in Helix aspersa that the developingembryo takes up calcium from the egg shell for the formationof its embryonic shell. However, other results from behaviouralobservations and calcium measurements in various reproductivestructures do not support the calcium hypothesis. We found thatthe dart penetrated the skin in 91.7% or the shootings, butit was internalized by the recipient in only 6.3% of the shootings.The amount of calcium in one dart is roughly equal to that ofone egg, and thus it would not contribute significantly to anaverage clutch of 59 eggs. The spermatophore contains virtuallyno calcium, and therefore it is unlikely that the dart signalsa donation of calcium with the sperm. The dart is also unlikelyto influence egg laying since dart shooting does not predicteither the latency or the productivity of egg laying in theshooter or the recipient. We conclude that the love dart ofHelix aspersa is not a gift of calcium. Instead, we suggestthat it is a vehicle to introduce a substance into the partnerto influence the fate of the donated sperm. (Received 8 November 1996; accepted 25 April 1997)     

NATURAL HISTORY AND POPULATION DYNAMICS OF THE LAND SNAIL HELIX TEXTA IN ISRAEL (PULMONATA: HELICIDAE)

Helix texta is endemic to the Mediterranean regions of Israel.It has a seasonal activity pattern which starts in the autumn,with the first rains, and dwindles towards the spring, whenthe snails dig into the ground for a six months long aestivation.A cold spell of 0°C will, however, terminate the activeseason of the adult snail, even in the middle of the rainy season.Survival of the young is very low and most of them (90%) donot survive their first year, because of the winter cold andthe summer drought. Massive predation of adult snails by wild boar was observedin December 1986: within a few days, about 50% of the adultsin the study in the area were eaten. A very rapid growth ofyoung and subadults was observed immediately after this predation.These observations suggest that the extent of recruitment ofnew adults to the population may be partly controlled by existingadults, through a growth-inhibiting pheromone in the mucus.After predation this inhibiting factor disappears, enablinga subsequent rapid growth of the young. In this manner, massive,irregular predation of the adults by a large predator, and changesin juvenile survival, result in sharp fluctuations in the agestructure of the population. The resulting pattern of unstablepopulation dynamics is different from that described for theEuropean species of Helix. (Received 16 January 1989; accepted 17 April 1989)     

FOOD SELECTION IN THE LAND SNAIL LIMICOLARIA FLAMMEA MULLER (PULMONATA: ACHATINIDAE)

Food consumption by juvenile and adult specimens Limicolariaflammea was measured as dry weight of food eaten. The snailwas fed weekly for 15 months on four types of food (lettuce,potato, apple and carrot) and chalk provided, hence an opportunityto select for food. Juvenile snails showed greater selectivitydepending on whether the food was needed for somatic growthor reproduction. On the other hand, when fed exclusively onone each of the same food items and given no opportunity tochoose, the snails grew better on the same food items they selectedin the choice experiment. The absorption efficiencies (AD) werehigh, the utilisation efficiency (ECI) was 1–13% and showeddistinct peaks corresponding to egg production. Values for ECDwere 1.25–15.5%. Variation in growth rates was due mainlyto utilisation efficiencies of the four foods tested. Largeamounts of chalk were eaten during the period of somatic growthand greatly reduced during the reproductive phase of L. flammea.The significance of these findings to the snails ecology isdiscussed. (Received 9 April 1991; accepted 11 June 1991)     

CALCIUM PROVISION TO EGGS IN TWO POPULATIONS OF HELIX ASPERSA BY PARENTS FED A DIET HIGH IN LEAD

The reproductive performance of two populations of Helix aspersawith different histories of exposure to lead was assessed bytheir egg production, hatching success and the metal contentof the eggs. By using laboratory-bred parents raised in a Pb-freeenvironment, the possibility of an inherited response to a highPb exposure was assessed. Following a period of dormancy, six individuals in each population werepaired for copulation and allowed to oviposit. During this time theywere fed a diet with 500 µg.g^-1 Pb, for a total of 14days. After oviposition, the egg mass from each pairing was weighed,counted and samples analysed. Metal concentrations in four tissuefractions of the parents (shell, hepatopancreas, reproductive systemand rest9) were also analysed. There was no significant transfer of Pb to the eggs in either population.There were differences in their egg massesthe population froman uncontaminated site, Rottingdean (RD), laid, on average,twice the number of eggs of the snails from Minera (MI), a contaminatedsite in North Wales. However the latter had a higher averagepercentage emergence (90% compared to 55% for RD). There wasalso considerable hatching asynchrony in the RD offspring, whilehatching within each MI egg mass was completed within two days.This is discussed in relation to the Ca status of the parentand possible differences in reproductive strategy between thetwo populations. (Received 22 February 2000; accepted 8 May 2000)     

NERVOUS CONTROL OF OVULATION AND EJACULATION IN HELIX ASPERSA

The ovotestis duct transports male and female gametes from theovotestis, through the seminal vesicle, and into the fertilizationpouch-spermathecal complex. All these structures are innervatedby small branches of the intestinal nerve. Electrical stimulationof the nerve increased the rate at which gametes were transportedin the duct and caused autosperm to flow into the fertilizationpouch-spermathecal complex. These events were accompanied bystimulation-induced peristaltic contractions along the ductand activation of the cilia lining the interior of the duct.Acetylcholine and serotonin were identified as excitatory transmittersin this system, while FMRFamide was identified as a muscle relaxant.The nervous control of ejaculation may contribute to optimizingthe size of the ejaculate in a context of sperm competition,while the involvement of the nervous system in ovulation maystem from a requirement for sensory integration that is peculiarto the Stylommatophora. (Received 16 December 2004; accepted 31 March 2005)     

GROWTH, MORTALITY, AND FEEDING RATES OF THE SNAIL HELIX ASPERSA AT DIFFERENT POPULATION DENSITIES IN THE LABORATORY, AND THE DEPRESSION OF ACTIVITY OF HELICID SNAILS BY OTHER INDIVIDUALS, OR THEIR MUCUS

When reared at high densities, young Helix aspersa show lessshell growth, even if waste products are removed. They alsofeed less, and show increased mortality. It is suggested thatthese effects are linked to reduced activity. Juveniles showreduced activity in the presence of adults or their mucus. Mucusof adult Cepaca nemoralis also depresses the activity of bothadult and young Helix and Cepaea. ^* Present Address: Unit Zoologi, Universiti Kebangsaan Malaysia,Jalan Pantai Baru, Kuala Lumpur, West Malaysia. (Received 12 May 1981;     

LAND USE AND SUMMER BIRD POPULATIONS IN NORTHWESTERN GALICIA, SPAIN

Summer bird populations were studied in ten sites: two heathlands, two maritime pine Pinus pinaster plantations, one, four, seven and 13-year-old eucalypt Eucalyptus globulus plantations, one climax oak Quercus robur woodland, and a farmland area in northwestern Galicia, Spain. The farmland and oak wood contained the greatest number of species (24 and 17 respectively) while the one and four-year-old eucalypt stands contained the fewest species (four and six respectively). Avian density was highest in the oak wood (93 birds/ 10 ha) and lowest in a heathland site, the four and one-year-old eucalypt stands (26,13, and seven bird/10 ha respectively). The wren was the most widespread and dominant species occurring in all study areas except the one-year-old eucalypt stand. Mean bird species diversity for the four eucalypt stands was significantly lower than mean bird species diversity for the other wooded habitats. The avian communities of the heathlands, maritime pine and eucalypt plantations, in contrast to the oak wood and farmland, show an excessively high dominance of one species. Using avian dominance as a biological index of habitat perturbation suggests that over half the landscape (4000 km²) in northwestern Galicia is suffering from chronic stress. The newest stress to the avifauna in a long history of land exploitation by man is the importation and cultivation on grand scale of eucalypts. Balancing this tendency, traditional, small-scale farming on the mountainous terrain leads to the development of many tiny parcels whose average size is 0.28 ha; this in turn creates much ‘edge’ in hedgerow which serves to maintain a relatively rich avian species diversity in farmland.     

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)     

CHOLINE ACETYLTRANSFERASE, ACETYLCHOLINESTERASE AND AROMATIC l-AMINO ACID DECARBOXYLASE IN SINGLE IDENTIFIED NERVE CELL BODIES FROM SNAIL HELIX ASPERSA

—The distribution of choline acetyltransferase, aromatic l -amino acid decarboxylase and acetylcholinesterase in the nervous system of Helix aspersa has been studied using homogenates of whole ganglia, microdissection from freeze-dried sections and dissection of single neurons from fresh tissue. Choline acetyltransferase was found in both the cell body and neuropil layers of all the Helix ganglia. The enzyme was not specifically localized to any ganglion or region of ganglion. Between 10 and 30 per cent of the isolated single cell bodies contained the enzyme. The enzymic activity corresponded to 50–200 mmol ACh/1 cell bodies/h. Choline acetyltransferase is probably a specific marker for cholinergic cells in this species. Aromatic l -amino acid decarboxylase was more selectivity localized and its distribution corresponded well with that of monoamine containing cells as visualized by the fluorescence histochemical technique. A large proportion of cell bodies were localized in the boundary between the visceral and right parietal ganglia and in the pedal ganglion. The other ganglia contained few such cells. The activity of aromatic l -amino acid decarboxylase corresponded 10–50 mmol dopamine/1 cell bodies/h. A method was developed to measure the enzyme activity towards 5-hydroxytryptophan and DOPA in single cells simultaneously. The ratio between the activity towards both substrates did not vary significantly for the different cells. The enzyme is probably a specific marker for monoamine cells, but cannot be used to differentiate between the different monoamine cells. Acetylcholinesterase was uniformly distributed in the ganglia and was probably present in all nerve cells.     

THE REPRODUCTIVE BIOLOGY OF THE AMPULLARIID SNAIL POMACEA URCEUS (MULLER)

Investigations of the reproductive biology, life cycle and populationdynamics have been undertaken to assess the potential of Pomaceaurceus as a culture species in Trinidad. The species is dioeciousand evidence from microscopic gonad analyses, together withmaturity indices and size frequency distributions in the population,all indicate that the reproductive cycle is annual. Adults spawnat the end of the rainy season months. Hatching and early developmentof the young occur in the dry season (January to May) whileadult females aestivate. The total developmental period variedbetween 22 to 30 days. Mean fecundity is 54 eggs/female witha range of 21 to 93 for egg masses deposited in the field andthose produced under laboratory conditions. ^*Present address: Zoology Dept., University of Aberdeen, TillydroneAve., Aberdeen AB9 2TN. Scotland. (Received 13 January 1988; accepted 11 April 1988)