COPULATION AS FEMALES AND USE OF ALLOSPERM IN THE FRESHWATER SNAIL GENUS BULINUS (GASTROPODA:PLANORBIDAE)

Each of 8 snails in 2 groups of Bulinus (Physopsis) globosus,1 group raised in isolation and 1 group raised in community,were paired for 14 consecutive days with a male-acting partnersnail. In each group, the experimental snails, which were notallowed to act as males, were able to copulate as females onapproximately 94% of the days paired. Two copulations as female,with the same male partner, occurred on 50% of the days thatthe snails were paired, in the 2 groups combined. Non-receptivefemale behaviour by the experimental snails occurred frequently,and copulation was prevented by such behaviour during 6 pairings,3 in each group. Young B. (P.) africanus first copulated as females when theywere 31–33 days old. The accessory sex glands of the femalereproductive tracts of these young female-acting snails containedmoderate to large amounts of secretion. B. (P.) africanus, which were raised in pairs, laidcross-fertilizedeggs in isolation for an average of 76 days, and 1559 eggs/snailwere deposited before cross-fertilization ceased. Cross-fertilizedeggs were produced for as long as 120 days. After 1 copulation as female, virgin B. (P.) africanus laidcross-fertilized eggs for an average of 78 days and deposited3654 eggs/snail before crossfertilization ceased. Cross-fertilizedeggs were produced for as long as 113 days. After 2 copulationsas female, 1 copulation on each of 2 consecutive days, virginB. (P.) africanus laid cross-fertilized eggs for an averageof 102 days and produced 4397 eggs/snail before cross-fertilizationceased. Cross-fertilized eggs were produced for as long as 123days. Snails which were homozygous for an allele governing mantlepigment pattern were raised with a partner which was homozygousfor a different pigment pattern. Young produced in a 4-day periodafter the snails were isolated were 100% heterozygous. The snailswere then rearranged into pairs with a partner of the same genotypefor 4 days, during which time 26% of the young produced werehomozygous. The snails were again isolated for 4 days, and 49%of the young produced during this 4-day period were homozygous.The results of this experiment strongly suggest that multipleoutcrossing occurred. In B. (P.) africanus, stored allosperm were used to fertilizeeggs after 1, 4 and 7 weeks of starvation; after 1 and 4 weeksof 15°C low temperature and 4 weeks of 15°C + 4 weeksof 10°C low temperature; and after 1 and 4 weeks of desiccation.After 8 weeks of desiccation, 2 of 3 surviving snails reproducedby self-fertilization and 1 snail did not reproduce. Too fewsnails survived 8 weeks of desiccation for a conclusion to bereached on the ability of allosperm to survive. (Received 1 June 1984;     

PATTERNS OF SEXUAL ROLES ADOPTED BY THE SCHISTOSOME-VECTOR SNAIL BIOMPHALARIA GLABRATA (PLANORBIDAE)

Relatively little is known about the mating behaviour of hermaphroditefreshwater snails, many of which transmit the Schistosoma trematodesamong humans in developing countries. Knowledge of the breedingbiology of these snails could help in the design of schistosomecontrol programmes, as well as possibly contributing to ourunderstanding of the evolution of simultaneous hermaphroditismin animaL. Here we describe an experiment investigating thepatterns of sexual roles adopted by the Schistosoma mansoni-vectotsnail, Btmphalaria glabrata. During observations on groups offreely interacting snails, no individuals copulated significantlymore often in the male than in the female role, or vice versa.Only one individual showed a pattern of alternating sexual rolesover successive matings that differed significantly from a randomsequence of roles. There was no evidence for reciprocal copulationwith one particular partner, either between consecutive matings(unless they were temporarily isolated from other snails) orbetween non-consecutive matings (separated by copulations withother conspccifics). We discuss these results in the contextof sex allocation and ESS mating strategy theories. ^*Present address (or correspondence 1 G VERNON. Bioscan (UK).Standingford House, Cave Street, St. Clements, Oxford OX4 IBA. (Received 5 October 1995; accepted 6 November 1995)     

The Neuropeptide APGWamide as a Penis Morphogenic Factor (PMF) in Gastropod Mollusks

The goal of this study was to further investigate the role ofendogenous APGWamide levels in imposex induction in snails.APGWamide is a common neurotransmittor/neuromodulator peptidefound in many species of molluscs, and is often related to sexorgan growth or reproductive behavior. Mud snails (Ilyanassaobsoleta) were collected from the Rachel Carson Estuarine Reservenear Beaufort, NC, and were dosed with the environmental contaminanttributyltin (TBT), testosterone (T), or solvent vehicle (EtOH/saline) controls. Both TBT and T have been shown previouslyto induce female snails to grow penises (a condition termedimposex), and to increase male penis size. Male normalized penislength was correlated to endogenous APGWamide levels (as measuredby Western blotting of whole animal homogenates, r² = 0.475),and control males had significantly higher APGWamide levelsthan control females. All TBT-treated animals, (male, female,and imposex) had levels of APGWamide similar to control malesand significantly higher than control females. In testosteronetreated animals, APGWamide levels were the same as controlsand it is likely that testosterone interferes with a downstreamsignaling event to induce imposex. In addition, immunohistochemistryfor APGWamide expression in abdominal areas was done on female,male and imposex snails collected from the wild. The patternof APGWamide in imposex snails was similar to male snails, showinglarge patches of immuno-reactive areas in the top portion ofthe visceral mass. In female snails, no areas of cross-reactivitywere found.     

THE COURTSHIP, PSEUDO-COPULATION BEHAVIOUR AND SPERMATOPHORE OF TURRITELLA COMMUNIS RISSO 1826 (PROSOBRANCHIA: TURRITELLIDAE)

Underwater field observations of reproductive behaviour in Turritellacommunis Risso were made in Killary Harbour on the west coastof Ireland during May and June in both 1986 and 1987. Thesesightings are documented and complemented by temperature, salinityand oxygen measurements, population structure, sex-ratio REMOTS®sediment profile imagery data and additional laboratory experimentationand observations. The population had a density of 639.5 individuals/m²at the study site with males comprising between 62.3 to 65.6%of the sexed population. Females were larger than males, thepopulation frequency distribution was negatively skewed andcharacterised by slow-growing, mainly adult individuals. Pseudo-copulationgroupings of 3 to 11 male specimens in star-shaped arrangementsappeared on the sediment surface in May and June as early summerbottom water temperatures increased above 10.0°C to 11.5°C.At its highpoint, c. 6 groups/m² were observed, with a meanof 5.9 males/grouping. Surface trails left by fertile malesindicated that they probably responded to a pheromone releasedby a passive fertile female lying within a radius of at least0.5 m. Smaller males were proportionately more active than largermales in sperm transfer. It was estimated that c. 15% of malesand 3% of females were synchronously fertile at the maximumobserved level of pseudocopulation activity. Laboratory observationsidentified the production of a complex doublewalled cylindricalspermatophore containing paired and unpaired euspermatozoa,and multiflagellate paraspermatozoa. Paired euspermatozoa weremost dominant and were arranged spirally at a 45° angleto the spermatophore axis. These were also frequently foundin morulae. An additional, spirallybound membranous tube exhibitingbulbous diverticula within the spermatophore contained paraspermatozoaonly. The observations confirm that sperm transfer is not apassive process as previously thought, and that the specieshas a far greater capacity for movement than was believed. Gregariousness,a noted feature among populations of turritellid species appearsto be a critically essential life strategy among these aphallic,internally-fertilising gastropods. Present address: Dr. J.J. Kennedy, Kantstrasse 38, 22089 Hamburg,Germany (Received 7 November 1994; accepted 15 March 1995)     

SIZE-ASSORTATIVE MATING IN A NATURAL POPULATION OF VIVIPARUS ATER (GASTROPODA: PROSOBRANCHIA) IN LAKE ZURICH, SWITZERLAND

The number of mating pairs, the size of the mating partners,and the distribution of individuals of Vivi-parus ater on agrid in Lake Zürich were recorded during one breeding seasonin 1990. There was positive assortative mating with respectto shell size. The proportion of copulating individuals rangedfrom 1% to 6% (average 3%) of the active population at any onetime. Individual snails copulated 60 times on average from Apriluntil November. Snails were abundant and copulated in shallowwater close to the shore in Spring. They moved towards deeperareas in Autumn. V. ater copulated on all substrates at anydepth (1–9 m)of the grid. The spatial distribution ofcopulations throughout the summer reflected the pattern of snailabundance. (Received 29 January 1993; accepted 14 November 1994)     

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)     

Shell-Boring Mechanism of the Gastropod, Purpura (Thais) lapillus: a Physiological Demonstration of the Role of Carbonic Anhydrase in the Dissolution of CaCO3

Measurement of carbonic anhydrase (CA) activity in homogenatesof accessory boring organs of the muricid gastropod,Purpuralapillus, by Meldrum and Roughton's manometric method showedthat CA is always present in boring as well as in inactive ABOs,but in variable amounts. Tests by the same method on whole ABOsin an isotonic solution were negative, proving that CA remainsintracellularly. Experiments in vivo on inhibition and activation demonstratedclearly that CA is responsible for demineralization of the valvesof lamellibranchs by Purpura:(a) at low concentrations of Diamox,partial inhibition of the enzymeoccurred, that is, the numberof complete holes decreased or disappeared and etchings increased;and at higher concentrations full inhibition took place. Inhibitionis reversed when snails are replaced in normal sea water; (b)action of pure CO₂ or mixtures of CO₂ and O₂ accelerated boring:in the optimal mixture, three times more boreholes were producedby snails than in the controls, and in about half the time.Under these conditions the reaction catalyzed by CA goes tothe right with hydration of CO₂ and is accompanied by releaseof H⁺ ions; in the presence of an increased concentration ofCO₂, the reaction is intensified and results in an additionalrelease of H⁺ ions. Consequently, destruction of CaCO₃ by theABO of Purpura in sea water enriched with CO₂ is accelerated. To identify the nature of the exchanged ion, we tested the effectof NaCl and KC1 on the boring mechanism in vivo by increasingthe amount of these salts in the seawater of snails with prey.In this case augmentation of the boring activity was noticed.These results suggest that the boring activity is accompaniedby complex ionic exchanges.     

GROWTH OF JUVENILE BITHYNIA TENTACULATA (PROSOBRANCHIA, BITHYNIIDAE) UNDER DIFFERENT FOOD REGIMES: A LONG-TERM LABORATORY STUDY

The growth of juvenile Bithynia tentaculata (Proso-branchia,Bithyniidae) was measured under controlled laboratory conditionsover 18 months. The animals were fed with different concentrationsof suspended food (Chlamydomonas reinhardii at 2, 000 cells.ml^–1and at 10, 000 cells.ml^–1), solid food (lettuce) and combinationsof both (lettuce with 2, 000 cells.ml^–1 and with 10, 000cells.ml^–1 Chlamydomonas). The growth of all animals wasmeasured weekly, and after 1 years final shell sizes, shelldry weights and ash-free dry weights, as well as soft body dryweights and ash-free dry weights were determined. Survival rateof the animals increased from 20% when fed with 2, 000 algalcells.ml^–1 to 75% with lettuce and 10, 000 cells.ml^–1as food. Final shell sizes and shell weights were not influencedby food, but soft body dry weights were significantly reducedwhen animals were fed on suspended food only. Reproduction (i.e.egg laying) was observed at the end of the second summer whenlettuce (with or without algal addition) was given as food.The ecological consequences of these results are discussed. (Received 7 April 1993; accepted 9 August 1994)     

DISTRIBUTION AND PRODUCTION OF THREE HYDROBIA SPECIES (GASTROPODA: HYDROBIIDAE) IN A SHALLOW COASTAL LAGOON IN THE BAY OF CADIZ, SPAIN

The abundance, life span, growth and production of the mud snailsHydrobia minoricensis, H. ulvae and H. ventrosa in a semi-naturallagoon system were studied by taking monthly samples at threesites during 1991 and 1992. The most abundant species, H. minoricensisoccurred at mean densities of 12834 to 26264 snails m^–2(10.7 to 25.8g dry weigh m^–2), depending on the site.The least abundant species, H. ulvae, occurred at mean densitiesof 185 to 353 snails m^–2 (3.2 to 2.2g dry weight m^–2).The numerical abundance and biomass of the three Hydrobia specieswere positively related to the biomass of benthic macroalgae(P<0.01). Although H. ulvae egg capsules were recorded throughoutthe year, newly hatched snailsof this species were not observed,in contrast to the other two species. The early spring and summercohorts of H. minoricensis and H. ventrosa seemed to be themost numerous. The average life spans of these two species wereestimated to be about 18 and 13 months respectively. Annualproduction estimates for the whole lagoon system were 29.0 (6.3),5.5 (0.8) and 5.2 (1.0)g dry weight (ash-free dry weight) m^–2yr^–1 for H. minoricensis, H. ulvae and H. ventrosa respectively.The annual P/B ratio was about 2 for H. minoricensis and H.ventrosa. (Received 5 July 1994; accepted 5 October 1994)     

Tales of two snails: sexual selection and sexual conflict in Lymnaea stagnalis and Helix aspersa

Sexual selection and sexual conflict have been shown to playkey roles in the evolution of species with separate sexes. Experimentalevidence is accumulating that this is also true for simultaneoushermaphrodites. For example, many species of land snails forcefullystab their mating partners with love darts. In the brown gardensnail (Helix aspersa, now called Cantareus asperses), this dartincreases sperm storage and paternity, probably via the transferof an allohormone that inhibits sperm digestion. A recent interspeciescomparison of dart-possessing land snails revealed coevolutionbetween darts and spermatophore-receiving organs that is consistentwith counteradaptation against an allohormonal manipulation.The great pond snail (Lymnaea stagnalis) seems to use a seminalproduct to manipulate its partner and mates in the male rolewhen enough seminal fluid is available in the prostate gland.Receipt of semen not only initiates egg laying in virgin animals,but also feminizes the mating partner later in life. These increasesin the female function have been shown to go at the expenseof growth and seminal fluid production of the sperm recipient.Although in Helix, and probably also Lymnaea, the sperm donorbenefits from the induced changes through increased fertilizationsuccess, the sperm recipient may experience injury, imposedreallocation of resources, and altered sperm storage. Thesefindings support the existence of sexual conflict in simultaneouslyhermaphroditic snails, and its importance for the evolutionof mating behaviors and reproductive morphologies is discussed.     

GROWTH AND REPRODUCTION OF THE MINUTE LAND SNAIL PUNCTUM PYGMAEUM (DRAPARNAUD)

Reproductive characters and growth of Punctum pygmaeum wereinvestigated under laboratory conditions. Eggs were laid singlyat intervals of 2–8 days. Egg size averaged 0.41 by 0.50mm. Due to a variable period of egg retention, newly depositedeggs contained differently developed embryos resulting in anincubation period varying from 1 to 34 days. Each snail on averagelaid 6.0 eggs (range 1–16 eggs) during its life. Egg productionrate and hatching success in individuals kept in isolation werethe same as in individuals kept in pairs. Hatchlings measured0.49 mm in shell breadth. Snails grew throughout their life,but the growth rate declined significantly at a shell breadthof 1.3–1.4 mm, the point at which the onset of sexualmaturity occurred. Comparison of reproductive characters ofsnails living in leaf litter revealed that very small-sizedspecies produce few, but in relation to their shell size largeeggs, which they oviposit singly. ^*Present address: Zoological Institute, University of Basel,Rhein-sprung 9. CH-4051 Basel, Switzertand (Received 23 May 1988; accepted 11 July 1988)     

安徽虫瘟霉的黍米培养及其对桃蚜的侵染力

安徽虫瘟霉Zoophthora anhuiensis (Li) Humber是较难人工培养的蚜虫专化性病原真菌。将灭菌并适度熟化的黍米Panicum miliaceum L.作为固体基质与挑碎的安徽虫瘟霉平板菌落混合,在20℃和12L∶12D的温光条件下静止固体培养,获得了产孢潜能大、杀蚜活性强的米粒培养物。培养7天的黍米的产孢量达13.0×10⁴个孢子/粒,产孢持续时间长达6天。用此黍米培养物弹射的孢子对桃蚜Myzus persicae (Sulzer) 若蚜进行7.9～134.9个孢子/mm²共9个剂量的孢子浴接种,所获数据很好拟合时间 剂量 死亡率模型。接种后第5～7天各天的LC₅₀依次为59.8, 39.5和33.5个孢子/mm²,LC₉₀依次为354,234和198个孢子/mm²。在57.7～134.9个孢子/mm²的接种剂量范围内,致死中时LT₅₀从5.1天下降到4.3天。由此表明,安徽虫温霉的黍米培养不仅简单易行,而且菌种的产孢和侵染生物学特性在培养物中被充分体现,每颗米粒如同自然罹病而死的蚜尸,值得进一步研究开发和利用。     

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;     

Role of the petasma and appendices masculinae during copulation and insemination in the penaeoid shrimp,Sicyonia dorsalis (Crustacea: Decapoda: Dendrobranchiata)

Summary

The role of presumed male gonopods in the insemination of females was investigated in the penaeoid shrimp Sicyonia dorsalis. Males with partially ablated petasmata or with ablated appendices masculinae did not copulate with females, while control males with exopods removed from the third pleopods copulated frequently and successfully inseminated females. However, males with petasmata altered by blockages at the tip or at the base did copulate with females but were unable to inseminate them. The hypothesis that the petasma of Sicyonia is a sperm injection device is rejected. Evidence against this hypothesis includes the structure of the gonopods, the position of the male at right angles below the female during copulation, the brevity of copulation, the frequency of copulation during normal matings, and the ability of males to inseminate only the spermatheca on one side per successful copulation. It is hypothesized that the male uses the petasma, supported by the endopods of the second pleopods, to hook onto the female thelycum, adjusting position so that one erect genital papilla directly injects sperm mass into the aperture of one of the paired spermathecae. It is suggested that female selection on male petasma structure and use has resulted in the complex form of the petasma, rather than a purely mechanical selection for an efficient sperm injection device.     

Effect of metanotal secretion ingestion on oviposition in a tree cricket, Truljalia hibinonis (Orthoptera: Gryllidae)

The female Truljalia hibinonis ingests metanotal secretions of the male during copulation. The effect of ingestion on oviposition behavior was compared between three female groups: females that copulated once with an intact male (a male that had not been manipulated; M group); females that copulated once with a male from which most of the metanotal secretion had been removed (NO group); and females that copulated once with an intact male followed by being artificially supplied with metanotal secretion three times (MS group). There were no obvious differences in female fecundity across the three groups. However, within the MS group, intake of an optimal amount of metanotal secretion increased the number of eggs laid. This effect appeared quickly after ingestion and was most effective on the first bout (eggs laid during the first few days after copulation) after ingestion of the metanotal secretion. In contrast, the number of eggs laid had a negative correlation with the amount of metanotal secretion ingested when the amount exceeded the optimal in this experimental arrangement.     

Vibrio cholerae ACE stimulates Ca(2+)-dependent Cl(-)/HCO(3)(-) secretion in T84 cells in vitro

ACE, accessory cholera enterotoxin, the thirdenterotoxin in Vibrio cholerae, has been reported toincrease short-circuit current (I_sc) in rabbitileum and to cause fluid secretion in ligated rabbit ileal loops. Westudied the ACE-induced change in I_sc andpotential difference (PD) in T84 monolayers mounted in modified Ussingchambers, an in vitro model of a Cl secretory cell. ACEadded to the apical surface alone stimulated a rapid increase inI_sc and PD that was concentration dependent andimmediately reversed when the toxin was removed. Ion replacement studies established that the current was dependent on Cland HCO₃. ACE acted synergistically with theCa²⁺-dependent acetylcholine analog, carbachol, tostimulate secretion in T84 monolayers. In contrast, the secretoryresponse to cAMP or cGMP agonists was not enhanced by ACE. TheACE-stimulated secretion was dependent on extracellular andintracellular Ca²⁺ but was not associated with an increasein intracellular cyclic nucleotides. We conclude that the mechanism ofsecretion by ACE involves Ca²⁺ as a second messenger andthat this toxin stimulates a novel Ca²⁺-dependent synergy.

     

Relative Importance of Ion Exclusion, Secretion and Accumulation in Spartina alterniflora Loisel.

The extent to which Spartina alterniflora Loisel. excluded,secreted or accumulated the major seawater ions (Cl^-, SO^2-₄,Na⁺, K⁺, Mg²⁺, and Ca²⁺) was investigated under varying salinitytreatments. From a quantitative viewpoint, ion exclusion wasmost prominent and accounted for 91–97% of the theoreticalmaximum ion uptake as a result of transpiration and growth.Of those ions taken up, approximately half was secreted fromthe shoots. Relative to K⁺, a disproportionate amount of Na⁺was excluded at the roots and secreted by the shoots. The concentrationwithin the tissues of S. alterniflora did not change with salinitytreatment for the majority of the ions examined, but Na⁺ wasmore than twice as concentrated at 40 g dm^-3 than at lOgdm^-3.Calculations of the flux of ions from salt marsh sediments tothe flood water via shoot secretion or stem/leaf turnover indicatethat these processes may be important to the ecology of S. alternifloraas mechanisms that limit the accumulation of salt within theroot zone.     

Membrane Depolarization by Hexacyanoferrate (III), Hexabromoiridate (IV) and Hexachloroiridate (IV)

Three artificial electron acceptors of different E_o and charge,hexacyanoferrate (III) (K₃Fe(CN)₆), hexachloroiridate (IV) (K₂IrCl₆),and hexabromoiridate (IV) (K₂IrBr₆), were compared with respectto their rate of reduction by roots of Zea mays L., the concomitantproton secretion, and to the effect on plasmalemma depolarization. It has been shown that these plasma membrane impermeable electronacceptors were reduced by a plasmalemma reductase activity.At low concentrations proton secretion was slightly inhibited,at higher concentrations, however, the rate of proton secretionwas stimulated. The root cell plasmalemma showed a transientdepolarization after addition of all three electron acceptors.The depolarization was concentration-dependent for the iridatecomplexes but not for hexacyanoferrate (III). For both iridatecomplexes maximum depolarization was reached at 50 µmoldm^–3. A hypothetical model as an explanation of the redox dependentproton secretion will be given. Key words: Hexachloroiridate (IV), hexabromoiridate (IV), hexacyanoferrate (III), plasmalemma redox, membrane potential, Zea mays     

UMBONIUM VESTIARIUM (L.) (GASTROPODA, TROCHACEA) AS THE FOOD SOURCE FOR NATICID GASTROPODS AND A STARFISH ON A MALAYSIAN SANDY SHORE

Umbonium vestiarium (L.) forms virtually the entire diet of3 (possibly 4) species of naticid snails and the starfish Astropectenvappa Mueller annd Troschel on some north Penang sandy shores.Umbonium comprises about 99% of numbers and tissue of macrofauna.Predation totalled some 1.75 Umbonium (ca. 33 mg dry tissue).m^-2.day^-1 across much of the downshore sand flats rising to 2.3Umbonium (ca. 45 mg). m^-2. day^-1 near MLWS. Natica maculosaLamarck comprised > 80% of the predators and took 77–94%of the Umbonium eaten. Natica antonii Phillippi alone addedto this toll on the upper reaches of the zone while Polinicesspp and Astropecten appear to have taken 12–14% of thetotal toll of Umbonium near MLWS. Total predation is indicatedat 237–327 kJ.m^-2. year^-1 across the shore and this representsalmost the total flow of energy from primary consumers to intertidalbenthic predators on such shores and accounts for some 15.6%(lower shore)—20.5% (upper shore) of total Umbonium production. (Received 10 September 1982;     

THE GROWTH AND BREEDING CYCLE OF LITTORINA OBTUSATA (GASTROPODA : PROSOBRANCHIATA) FROM CARDIGAN BAY

The population structure of Littorina obtusata was for mostof the year bimodal, with a stable adult component and an activelygrowing juvenile component. Growth rate of the juvenile componentwas similar at different beach levels and there was a retardationof growth as the snails approached sexual maturity. There was little variation in the histology and external appearanceof the reproductive organs throughout the year and the breedingcycle has been characterized by the amount of spawn laid andthe presence of mature eggs in the oviduct of the female. ^*Present address: Portway School, Penpole Lane, Shirehampton,Bristol. (Received 7 September 1977;