THE ROLE OF LYMNAEA AURICULARIA (LINNAEUS) AND LYMNAEA PEREGRA (MULLER) (GASTROPODA: PULMONATA) IN THE TRANSMISSION OF LARVAL DIGENEANS IN THE LOWER THAMES VALLEY

Eleven species of larval digeneans are reported for the firsttime in the UK from the freshwater snail Lymnaea (Radix) auricularia.The latter replaced its congener L. peregra in a gravel pitnear Wraysbury, Berkshire in the Lower Thames Valley and hostedan assemblage of larval digeneans similar to that previouslytransmitted by L peregra. The larval digeneans of L. peregrain small bodies of water in the area were also similar to thosefound in L. auriculania in the gravel pit. This replacementin the role of the host is explained on the basis of morphological,physiological and ecological similarities of the two lymnaeidspecies. (Received 19 February 1992; accepted 27 April 1992)     

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;     

STARVATION, DESICCATION AND USE OF ALLOSPERM IN THE HERMAPHRODITE FRESHWATER SNAIL BIOMPHALARIA GLABRATA (GASTROPODA: PULMONATA)

Pairs of albino and pigmented snails were used to test the hypothesisthat the hermaphrodite freshwater gastropod Biomphalaria glabratawas able to conserve and use allosperm, despite periods of desiccationand starvation. In laboratory experiments, lots of twenty snailswere subject to 0, 5, 10, 15 AND 20 days starvation; similarlots of 20 snails were subjected to 0, 2, 4, 6, 8, 10 weeksof desiccation at R.H. 100%. After all periods of starvationand desiccation, albino parents were still producing significantnumbers of pigmented offspring, suggesting preferential cross-fertilizationusing stored allosperm. (Received 30 September 1993; accepted 23 December 1993)     

THE MORPHOLOGICAL EXPRESSION OF IMPOSEX IN NUCELLA LAPILLUS (LINNAEUS) (GASTROPODA: MURICIDAE)

The prosobranch gastropod Nucella lapillus exhibits imposex(occurrence of male parts in addition to the female genitalduct) in response to tributyltin (TBT) pollution. Six stageswith three different types in the first three stages and twotypes in the last two stages can be distinguished and are documentedwith SEM-photographs for the first time. Furthermore, nine additionalalterations to the genital tract are shown. As a consequence of TBT exposure in early life stages a sexchange might occur. The oogenesis is suppressed and supplantedby spermatogenesis. The results are two observed specimens witha male genital tract (with a testis) and different female parts.Anatomical and histological details are given here. If TBT-expositionoccurs later, the vagina is replaced by a small prostate glandor the vaginal opening is occluded by proliferating tissue. The two established indices for PSH monitoring (VDS and RPS)in muricid gastropods are discussed. It is shown that the VDSis the more valid parameter. (Received 2 January 1991; accepted 1 March 1991)     

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

The presence of connective interstitial cells in the wall ofseveralorgans of the distal reproductive system in Oxychilusatlanticusis described by means of scanning electron microscopy(SEM).Connective interstitial cells extend from the distal part ofthevas deferens to the penis, being the dominant cell type of largeportionsof the epiphallus as well as in the whole penial caecum(=flagellum).These cells are highly mineralized and are characterizedbythe presence of large amounts of solid spherules (up to 4 mmindiameter) in their cytoplasm. A semi-quantitative evaluationby X-raymicroanalysis demonstrates that calcium and phosphorusare the mainelements of the spherules. Their function is unknown,but they couldaid in copulation or they could assist the survivalof the transferredspermatozoa. (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;     

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;     

SPERM AND SPERMIOGENIC ULTRASTRUCTURE IN THE MATHILDIDAE WITH A REVIEW OF SPERM MORPHOLOGY AND ITS SYSTEMATIC IMPORTANCE IN THE ARCHITECTONICOIDEA (GASTROPODA)

The ultrastructure of spermiogenesis and spermatozoa is examinedfor the first time in the heterogastropod family Mathildidae(Mathilda brevicula Bavay and Mathilda sp.). Mathildid spermatozoaexhibit the following features: (1) an ovoid acrosomal vesicleshowing curved dense layers basally and underlain by a thincurved plate; (2) a solid, rod-shaped nucleus showing a shallowbasal invagination (nucleus helically keeled in Mathilda sp.);(3) a midpiece composed of a 9+2 axoneme, 9 thick and periodicallybanded coarse fibres (associated with the axonemal doublets)and a continuous mitochondrial derivative; (5) glyco-gen piececomposed of the axoneme surrounded by dense granules; (6) andend piece (paddle-shaped terminally). Results of the study demonstratethat mathildids share the same key sperm and spermi-ogenic featuresobserved in the Architectonicidae. The monophyly of the Architectonicoidea,established by previous authors on anatomical grounds, is thereforeconfirmed using sperm morphology. Spermatozoa of the two mathildidsinvestigated lack the periodically banded midpiece structuresseen in investigated species of Architectonica (A. pcrspectiva(Linnaeus), A. perdix (Hinds)), Heliacus (H. (Heliacus) variegatus(Gmelin)) and Pseudotorinia (P. laseronorum (Iredale)) therebyresembling most closely the spermatozoaof Philippia lutea (Lamarck)and Psilaxis oxytropis (A. Adams) which also lack suchbandedstructures. Given the widely accepted primitive status of theMathildidae within the Architectonicoidea, the genera Philippiaand Psilaxis may prove to be relatively basal rather than advancedgenera within the Architectonicidae. Thi: view needs to be testedby examination of sperm morphology in other genera of the Architectonicidaeand also the basal mathildid genus Tuba sensu lato (= Gegania+ Tubena). Such information will be vital in any future cladisticstudy of the Architectonicoidea. (Received 16 June 1994; accepted 6 February 1995)     

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)     

THE ONCHIDIACEA (GASTROPODA, PULMONATA) OF HONG KONG WITH A WORLDWIDE REVIEW OF THE GENERA

The Onchidiacea are a superfamily of mainly marine intertidal,slug-like pulmonates. Twelve genera are recognizable, and theseare redefined. Four of these genera are represented in HongKong, each by a single species, and these are recorded herein.Two of these are new; a third has not been seen since it wasoriginally figured in 1850. ^*Present address: Palmers Sixth-Form College, Grays, Essex. (Received 18 August 1983;     

GEOGRAPHICAL PATTERNS OF GENETIC DIFFERENTIATION IN THE LANDSNAIL HELIX ASPERSA MuLLER (GASTROPODA: PULMONATA)

In order to investigate the initial processes responsible forthe geographical distribution of the Mediterranean landsnailHelix aspersa, populations from 64 locations in Maghreb andin Europe were studied at 17 enzyme loci by both starch andpolyacrylamide gel electrophoresis. Allele frequencies and genotypedistributions were analysed to provide information on populationstructure and genetic differentiation among populations. Valuesof Nei's unbiased (D) and Rogers (R) genetic distances led tothe formation of two main groups, namely Europe-west Algeriaand east Maghreb (average D = 0.182). Such splitting supportedthe pattern of anatomical variation recently described. Twosamples from Morocco were excluded from these groups becauseof an average D = 0.224. Within this Moroccan entity, geneticdivergences did not distinguish the ‘giant’ snailsH. a.maxima from the brownsnails H. a. aspersa. Such biochemicalsimilarities among morphological heterogeneity throws the subspecificstatus of H. a. maxima into question. Values of fixation indicesindicated that the greatest differentiation between samplesoccurred on the Maghreb territory ( = 0.393). The patterns of population differentiation are discussed inrelation to past geological and historical events. Thus, threescenarios, not mutually exclusive, are considered: the disjunctionof populations by plate tectonics during the Tertiary, the formationof geographically isolated populations during the Pleistoceneglacial/interglacial cycle and the possibility of human introductionas early as the Neolithic period. (Received 9 December 1992; accepted 11 November 1993)     

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)     

THE LIFE CYCLE, POPULATION DYNAMICS, GROWTH AND SECONDARY PRODUCTION OF THE SNAIL BRADYBAENA FRUTICUM (MULLER, 1774) (GASTROPODA PULMONATA) IN NORTHERN GREECE

The life cycle, population dynamics, growth and secondary productionof the pulmonate land snail Bradybaena fruticum were studiedin northern Greece. The demographic analysis of the populationsof B. fruticum revealed that a) two cohorts exist in the fieldthroughout the year b) the reproductive period started in thebeginning of summer and c) growth was increased during springand autumn. According to von Bertallanffy' method B. fruticum needs 5 yearsto attain its maximum size in the field of 25.40 mm. Mortalityrate increases and life expectancy decreases with increasingage. Net reproductive rate (Ro) was equal to 3. 15 and per capitarate of increase (r^c) was equal to 1. Annual secondary production, calculated by Hynes' size frequencymethod, revealed a mean annual density of 5.9 individuals/m²,a mean standing crop (B) of 8.09 g/m²/year and an annual production(P) of 1.92 ± 0.11g/m²/year. Annual turnover ratio (P/B)was equal to 2.37. (Received 25 May 1989; accepted 17 March 1989)     

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;     

WATER CONTENT OF SLUGS (GASTROPODA:PULMONATA) IN NATURAL HABITATS, AND THE INFLUENCE OF CULTURE CONDITIONS ON WATER CONTENT STABILITY IN ARION ATER (LINNE)

10 species of slug were collected from a variety of habitats.Within species, water contents were relatively constant, andit was concluded that they were maintained by water uptake,mainly through the integument, together with appropriate cryptozoicbehaviour. Weights of A. ater individuals held outdoors didnot show large daily fluctuations. ^*Present address: Department of Zoology, East Mailing ResearchStation, Maidstone, Kent ME19 6BJ (Received 15 June 1982;