AN ULTRASTRUCTURAL STUDY OF SPERMATOZOA OF HELIX ASPERSA AND HELIX POMATIA (GASTROPODA, PULMONATA)

Spermatozoa of the pulmonates Helix aspersa Müller andH. pomatia Linnaeus are examined in detail using transmissionelectron microscopy (TEM). Important features such as the acrosome,perinuclear sheath, nucleus and terminal region of the midpieceare described for the first time. Also presented are the firstultrastructural observations on spermatozoa from spermatophoresin any pulmonate gastropod (H. aspersa). No morphological differencescould be found between sperm taken from spermatophores and thosewithin the hermaphrodite duct in H. aspersa. Spermatozoa ofH. aspersa and H. pomatia snow all the characteristics of euthyneuranspermatozoa, namely: a helically-keeled nucleus; distinctivearrangement of acrosomal components (apical vesicle, acrosomalpedestal), and extremely elongate midpiece (axoneme and glycogenhelix enclosed by matrix and paracrystalline layers). The spermnucleus of both species is short, and the midpiece also formsthe terminal portion of the spermatozoon (glycogen piece absent).The extraordinary positioning of the acrosome in H. aspersa—reflectedbackwards from the nuclear apex—is not observed in H.pomatia, though a perinuclear sheath (possibly another acrosomalcomponent) is present in sperm of both species. Helix spermatozoaare compared with other euthyneuran sperm and briefly discussedfrom the systematic viewpoint. Present address: Department of Zoology, St. Lucia, 4067, Brisbane,OLD, Australia (Received 23 May 1988; accepted 17 August 1988)     

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)     

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)     

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;     

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;     

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)     

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)     

CALCIUM PHOSPHATE GRANULES IN THE REPRODUCTIVE SYSTEM OF OXYCHILUS ATLANTICUS (GASTROPODA: PULMONATA)

The presence of connective interstitial cells in the wall ofseveral organs of the distal reproductive system in Oxychilus atlanticusis described by means of scanning electron microscopy (SEM).Connective interstitial cells extend from the distal part of thevas deferens to the penis, being the dominant cell type of large portionsof the epiphallus as well as in the whole penial caecum (=flagellum).These cells are highly mineralized and are characterized bythe presence of large amounts of solid spherules (up to 4 mmin diameter) in their cytoplasm. A semi-quantitative evaluationby X-ray microanalysis demonstrates that calcium and phosphorusare the main elements of the spherules. Their function is unknown,but they could aid in copulation or they could assist the survivalof the transferred spermatozoa. (Received 7 June 1999; accepted 8 September 1999)     

THE STRUCTURE OF THE SECONDARY GILLS OF SIPHONARIA CAPENSIS (GASTROPODA: PULMONATA)

The secondary gill of the pulmonate limpet Siphonaria capensisis located in the dorsal portion of the mantle cavity. Eachlamella of the gill is triangular in shape and bears ciliarytufts which have a density of about 400/mm². The free ends ofsome of the cilia are enlarged biconcave discs. Each gill lamellais covered by a single layer of cuboidal (4x4 µm) epitheliumon each side, separated by a central haemocoelic space. Ciliatedand mucus secreting cells are interspersed amongst the epithelialcells. The haemocoelic space is spanned at intervals by trabeculaecontaining longitudinal and transverse muscle fibres. (Received 29 August 1986;     

A REVIEW OF LARVAL DEVELOPMENT IN THE INTERTIDAL LIMPET GENUS SIPHONARIA (GASTROPODA: PULMONATA)

A compilation of distributional and life-history data relatingto mode of larval development is presented for 26 species ofSiphonana, a genus of intertidal pulmonates. Most species depositgelatinous benthic egg masses with two species releasing pelagicegg masses. Fifteen species hatch as planktonic-developing larvae,nine hatch as direct-developing juveniles, and in a furthertwo larvae hatch with both the swimming velar apparatus (associatedwith plank-tonic development) and a crawling foot (associatedwith direct development). Data on mode of larval developmentare interpreted with respect to some adaptive models. Despiteimportant exceptions, there is support for adaptive models basedupon egg capsule size (direct developers hatch from larger eggcapsules) and intertidal distribution (direct developers generallyoccur higher on the shore than planktonic developers). Worldwide,planktonic developers are more widespread than direct-developingspecies, and individual planktonic species have a greater meanlatitudinal range. The evidence for adaptive models relatinglatitudinal distribution to developmental mode is equivocal.There appears to be no clear relationship between body sizeand developmental mode in the genus, although the smallest specieshas direct development and the largest has planktonic development.In most siphonariid subgenera, developmental mode appears tobe constant, but two subgenera contain a mixture of developmentaltypes (Received 1 November 1993; accepted 15 April 1994)     

WATER-CONTENT OF SLUGS (GASTROPODA:PULMONATA) MAINTAINED IN STANDARDISED CULTURE CONDITIONS

A culture method for mass-rearing slugs is described and thewater-contents of 11 species of slug raised under these conditionsare analysed. The conclusion is drawn that, supplied with anappropriate environment, slugs showed no obvious rhythmic fluctuationsin hydration but exhibited water-contents relatively stablewithin species and presumably maintained by behavioural mechanisms. ^*Present address: Department of Zoology, East Mailing ResearchStation, Maidstone, Kent ME19 6BJ (Received 12 May 1981;     

THE INFLUENCE OF HUMIDITY, PHOTOPERIOD, AND TEMPERATURE ON THE DORMANCY AND ACTIVITY OF HELIX LUCORUM L. (GASTROPODA, PULMONATA)

The dormancy of Helix lucorum from N. Greece is controlled primarilyby low humidity. Photoperiod and temperature have no directeffect, except in the anomalous combination of long cool daysor short hot days. Adults hibernate more readily than juveniles.The duration of hibernation (before rejection of the calcareousepiphragm) is only slightly shorter at high temperatures. (Received 15 December 1985;     

PATTERNS IN THE GEOGRAPHICAL RANGES OF SPECIES

1. A growing number of studies have documented patterns in species geographic ranges relevant to the study of community structure. These include patterns in the frequency of geographic ranges of different sizes, and in the interaction of range sizes with population abundances and variabilities, body sizes, trophic characteristics and extinction probabilities.
2. Agreement between hypothesized and observed relationships is reasonably good, but we do not know how general the patterns are. Analysis of patterns has focused upon a few taxonomic groups, a bias largely resulting from a lack of information on the geographic distributions of most taxa.
3. Many of the patterns are interrelated, and although theoretical bases to all the patterns can be suggested, it is possible that some are artifacts.
4. Taylor power plots give us a means of making predictions about population behaviour as it pertains to geographic ranges. Some of these predictions suggest that previous conceptions of such interactions have been too narrow, but empirical analyses of these patterns will be hampered by the difficulty of measuring population variability.
5. In general, our knowledge of the structure, and spatial and temporal behaviour, of species geographic ranges remains poor.     

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)     

ASPECTS OF THE LIFE CYCLE, POPULATION DYNAMICS,GROWTH AND SECONDARY PRODUCTION OF THE EDIBLE SNAIL HELIX LUCORUM LINNAEUS, 1758 (GASTROPODA, PULMONATA) IN GREECE

This paper examines the biology and ecology of Helix lucorumL. which lives in mainland Greece, as well as its growth andsecondary production. A demographic study revealed that (a)3 cohorts exist at any time during the year (when adults ofall generations belong to the same cohort) (b) egg-laying andhatching occur during the months of July and August respectively,(c) the most rapid growth takes place during spring. Study ofH. lucorum genitalia in relation to age showed that the snailsaxe sexually mature 3 years after hatching, when the largestdiameter of their shell (D) is equal to or greater than 35 mm Von Bertallanffy's method suggests that Helix lucorum may liveup to 14 years or more in order to reach its possible maximumsize (48.80 mm) The study of relative growth of D in relation to P_s (peristomesurface) of Helix lucorum shows that D grows faster than P_swhen D12.50 mm; juveniles change their growth rate when theirD arrives at 22.05 mm, and growth becomes slower when adultsarrive at 36.27 mm Annual secondary production calculated by the size frequencymethod gave a mean annual density of 3.39 individuals per m²,a mean annual crop (biomass) of 4.04 g-m^–2 and an annualproduction (P) of 5.02 g · m^–2. The annual turnoverratio (P//b) is equal to 1.24 (Received 23 June 1987;     

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)     

NOTES ON THE INTRODUCED TERRESTRIAL PULMONATA (GASTROPODA:MOLLUSCA) OF NEW ZEALAND

The New Zealand Pulmonate fauna contains 22 naturalized terrestrialspecies. For each, a bibliography is given for the New Zealandliterature and comment made on its biology and distribution.Vertigo ovata SAY and Oxychilus draparnaldi (BECK) are newlyrecorded from New Zealand. (Received 11 February 1981;