PRELIMINARY INVESTIGATIONS OF THE ECOLOGY OF THE INTERTIDAL PREDATORY GASTROPOD LEPSIELLA VINOSA (LAMARCK) (GASTROPODA MURICIDAE)

Lepsiella vinosa\ (Lamarck), a common intertidal predatory gastropod,was studied in two zones on an exposed rocky shore in Victoria(Australia). Those from the Mussel Zone in the mid-shore primarilyate one species of mussel and had a faster rate of energy consumptionthan those from the Littorinid Zone in the high-shore, whichmainly ate one species of littorinid gastropod. Shell length,growth rate and size-specific body weight of L. vinosa wereall significantly greater in the Mussel Zone. Egg capsules ofL. vinosa from the Mussel Zone contained significantly moreeggs and more potential hatchlings per capsule than those fromthe Littorinid Zone although there were no significant differencesin the sizes of eggs or hatchlings between zones. These differencesbetween the two zones were correlated to the greater consumptionof energy by L. vinosa in the Mussel Zone and were discussedin relation to the absence of dispersive larvae in this species ¹Present address: Department of Zoology, Monash University,Clayton, Victoria, 3168, Australia ²Present address: School of Biological Sciences, Carslaw Building,F07, University of Sydney, N.S.W., 2006, Australia. Addresseefor reprints. (Received 2 June 1987;     

HYBRID FITNESS SEEMS NOT TO BE AN EXPLANATION FOR THE PARTIAL REPRODUCTIVE ISOLATION BETWEEN ECOTYPES OF GALICIAN LITTORINA SAXATILIS

Partial reproductive isolation between closely related groupsof organisms is suggested to be of central importance duringspeciation. Galician populations of Littorina saxatilis arephenotypically differentiated into an upper-shore and a lower-shoremorph. These mate assortatively in the mid-shore zone of overlap,and genetic assessment shows an impeded gene flow between theparental morphs. The traditional explanation as to why assortativemating occurs is that reproductive isolation is reinforced dueto hybrid unfitness. Earlier studies have, however, not foundhybrids to be less viable. Likewise, growth and migratory behavioursare merely intermediate between those of the parental morphs.In the present study we compared male and female fertilitycomponents of the parental morphs and the hybrids to test hypothesesof decreased hybrid fertility. The results showed that hybridmales were as fertile as other males, and hybrid females didnot produce fewer embryos, nor aborted embryos at a higher rate,than the parental morphs. ^* To whom correspondence should be addressed. (Received 19 March 1999; accepted 7 July 1999)     

EMBRYONIC ABNORMALITIES IN POPULATIONS OF LITTORINA SAXATILIS (OLIVI) (GASTROPODA: PROSOBRANCHIA) IN THE WHITE SEA

The incidence of abnormally developing embryos in brood pouchesof females was studied in five populations of Littorina saxatilisliving under different salinity conditions. The proportion ofabnormals decreased during embryonic development. Age of femaleand infection with the ciliate Protophrya ovi-cola had no effecton the frequency of abnormals. The proportion of abnormal eggs(PAE) increased at the end of the reproductive season; a similartrend was found for the proportion of abnormal veligers (PAV)although it was not always significant. The proportion of abnormaljuveniles (PAJ) showed no seasonality. The increase of PAE coincidedwith decrease of water temperature in September. Environmentalsalinity was significantly correlated with frequency of abnormalsin L. saxatilis populations: the highest values of PAV and PAJwere observed in estuarine population (salinity range 5–8%),intermediate in populations living under conditions of 10–20%osalinityand lowest in control populations (23–24%). Itis suggested that critical salinity (sensu Khle-bovich, 1974)(approximately 8–10%) may also be a crucial point forstability of embryonic development in the White Sea populationsof L. saxatilis. (Received 28 June 1994; accepted 28 November 1994)     

PATTERNS OF REPRODUCTION IN FOUR LITTORINA SPECIES

The reproductive periodicities of four Iittorinids, Littorinaneglecta, L. nigrolineata, L. saxatilis and L. arcana were examinedin sympatric populations at Porth Swtan, Anglesey, North Wales.Three species, L. neglecta, L. nigrolineata and L. arcana, werefound to show marked reproductive periodicity. L. saxatilisshowed no periodicity and large numbers of animals were foundin breeding condition throughout the year. The oviparous speciesL. nigrolineata and L. arcana, came into reproductive conditionin the late summer or early autumn, L. nigrolineata in Augustand L. arcana in September, and their populations at Porth Swtanhad few individuals in breeding condition in the late springand early summer. L. neglecta, however, showed a periodicitywith high numbers of breeding animals throughout the springand summer and very low numbers in the autumn. For L. neglectathe basis of reproductive periodicity is shown to be the synchronizedannual life cycle of individuals of this species. It is thereforebelieved to differ from the other three species in which individualsare longer-lived and are supposed to undergo seasonal periodsof reproductive inactivity. (Received 2 March 1982;     

POPULATION GENETICS OF A SNAIL SPECIES COMPLEX IN THE BRITISH ISLES: LITTORINA SAXATILIS (OLIVI), L. NEGLECTA BEAN AND L. TENEBROSA (MONTAGU), USING SSCP ANALYSIS OF CYTOCHROME-B GENE FRAGMENTS

The intertidal snail Littorina saxatilis displays a range ofshell morphologies associated with a variety of habitats. Sincemorphology has an environmental and genetic basis, shell-basedtaxonomy may not accurately reflect genetic relationships. Weexplored genetic structure among adjacent populations of L.saxatilis (the robust open-shore type), L. neglecta (the tinybarnacle-dwelling type) and L. tenebrosa (the fragile brackish-watertype), at nine sites in Britain. Using single-strand conformationalpolymorphism analysis of a 375bp fragment of cytochrome-b wefound no evidence of species distinction. In AMOVA tests significantvariation was contained among populations (68%) and among individuals(32%, both P < 0.001), and insignificant variation was foundamong ecotypes. Genetic patterns suggested gene flow among ecotypesover small scales and a strong random input over larger scales. ^* To whom correspondence should be addressed. (Received 27 March 2000; accepted 10 July 2000)     

Habitat-dependent Ecotype Micro-distribution at the Mid-shore in Natural Populations of <Emphasis Type="Italic">Littorina saxatilis</Emphasis>

On the NW coast of Spain, there is a hybrid zone in natural populations of L. saxatilis, where the largest ecotype is adapted to the desiccation and heat stress of the upper shore, whereas the smaller one is adapted to the wave impacts of the lower shore. The two ecotypes meet and hybridise in a mid-shore area producing intermediate individuals or hybrids. It has been postulated that this hybrid zone is maintained by a selection-gradient model that assumes habitat-dependent selection for the different environments of upper and lower shore areas. In this study, we focus on the hybrid (mid-shore) area, where a transition occurs between upper-shore and lower-shore environments, in order to ascertain whether the habitat-dependent selection is maintained at the micro-habitat scale. We present data on snail density at several levels of the mid-shore (varying accordingly in the relative frequency of mussels and barnacles) obtained for three seasons in three consecutive years and three localities. In the mid-shore, the abundance of the RB ecotype increases with increasing tidal height, whereas the abundance of the SU ecotype increases with decreasing tidal height, suggesting that habitat-dependent selection is maintaining the ecotype micro-distribution.     

ESTERASE VARIABILITY IN THE GASTROPODS LITTORINA SAXATILIS (OLIVI) AND L. ARCANA ELLIS

The esterase system has been studied using isoelectric focusingin two closely related rough periwinkles, Littorina saxatilisand L. arcana, in an investigation of possible differences betweenthe two species at this level of discrimination. Collectionsof both species were obtained from Yorkshire, north Cornwalland south Devon; L. saxatilis was also obtained from sites insouth Cornwall (in and near the estuary of the Tamar), Dorsetand Scotland, where L. arcana was not present. There is considerablevariation both within and between sites which tends to maskany overall differences between the two species. However, ingeneral there is a much greater degree of heterogeneity withinsamples of L. saxatilis than within samples of L: arcana whenboth species are found together; this is reflected in the ShannonWiener diversity index for bands. Also the number of esterasebands is greater for L. saxatilis than for L. arcana at eachof the five sites where both occurred. A dendrogram derivedfrom the percentage occurrence of each band present in eachsample produced a reasonable degree of separation of the twospecies (Received 8 January 1988; accepted 24 February 1988)     

THE GENETIC RELATIONSHIPS OF THREE TAXA IN THE LITTORINA SAXATILIS SPECIES COMPLEX (PROSOBRANCHIA: LITTORINIDAE)

Three species in the Littorina saxatilis complex (the ovoviviparousL. saxatilis and the two oviparous species L. arcana and L.nigrolineata) were screened for the products of up to 22 lociusing gel elec-trophoresis. Analysis of allele frequencies andderived genetic distances showed that, with rare exceptionsin L. saxatilis and L. arcana, conspecific populations clusteredtogether. Sympatric pairs of populations showed significantdifferentiation at many polymorphic loci. The results confirmedthe biological reality of the three taxa, and did not suggestthe existence of any previously unrecognized species. The speciesare genetically closely related to each other, with Nei geneticdistances ranging from 0.035 to 0.083. (Received 24 October 1989; accepted 18 April 1990)     

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;     

Sympatric speciation as a by-product of ecological adaptation in the Galician Littorina saxatilis hybrid zone

On exposed rocky shores in northwestern Spain there is a strikingpolymorphism of Littorina saxatilis that has been claimed asan example of a putative sympatric ecological speciation process.Two ecotypes of this species have evolved that are adapted todifferent shore levels and habitats, although they meet andhybridize on the mid shore where their two habitats overlap.As a by-product of adaptation these ecotypes have evolved anincomplete premating reproductive isolation where they meeton the mid shore. Although they are not yet true species, andthe final outcome of the process cannot be predicted, the ecologicalmechanisms responsible for this polymorphism could cause sympatricspeciation in similar situations. Here, I review all data insupport of these claims and discuss the interest of such a modelsystem in microevolutionary studies. (Received 13 February 2006; accepted 25 October 2006)     

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)     

Habitat scale, female cannibalism, and male reproductive success in three-spined sticklebacks (Gasterosteus aculeatus)

We studied the effects of habitat scale (pool size) on the riskof egg cannibalism and the breeding behavior of three-spinedsticklebacks (Gasterosteus aculeatus). Laboratory experimentsshowed that in pools with similar population density, a largerfraction of males nested in pools of 4.8-m² surface area thanin 15-m² pools. Breeding males in the 4.8-m² pools also hadbigger territories, obtained more eggs, and were more aggressivetoward intruders than those in the 15-m² pools. The numbersof raids attempted on nests were similar in the two pool sizes.However, raiding-shoal sizes were larger, and hence individualsin these larger shoals were more succesful at overwhelming maleterritorial defenses in the 15-m² pools than in the 4.8-m² pools.At the field site, a salt marsh containing hundreds of tidepools of different sizes, sticklebacks settled at similar densitiesin small (<30 m² surface area) and large (>355 m²) pools.This resulted in significantly more fish being present in largepools than in small pools, a concomitant increase in the amountof nest raiding, and a decrease in individual male reproductivesuccess in large compared to small pools. A photogrammetry analysisof the salt-marsh pools suggests that the majority of fish breedingat the site will be subjected to a high risk of cannibalismbecause most fish nest in large pools. We conclude that spatialscale has a major influence on this species' breeding behaviorand reproductive success.     

Seasonal variation of reproduction rates and body size of Calanus sinicus in the Southern Yellow Sea, China

Annual variations of egg production rate (EPR) and clutch sizeof Calanus sinicus, as well as body size of females (prosomelength and dry weight), were investigated at a series of stationsin the Southern Yellow Sea by onboard incubation. Calanus sinicuswas spawning in all the 11 cruises investigated, and the annualvariation of EPR was bimodal. Monthly average EPR was highestfrom May to July, respectively, 5.97, 5.36 and 6.30 eggs female^–1d^–1, then decreased dramatically to only 1.37 eggs female^–1d^–1 in August and attained the lowest 1.07 eggs female^–1d^–1 in October. In November, average EPR increased againto 4.31 eggs female^–1 d^–1. Seasonal variation ofclutch size was similar to EPR, except that it decreased graduallyafter August rather than dramatically as did EPR. Prosome lengthof females was maximum in May and minimum in October, but dryweight was highest in November. Monthly average EPR correlatedbetter with prosome length than dry weight, while clutch sizewas rather determined by dry weight of females. It is suggestedthat egg production of C. sinicus was active during two discontinuousperiods when both surface and bottom temperature fell into itsfavorite range (i.e. 10–23°C), and different reproductivestrategies were adopted in these two reproductive peaks: otherthan the highest EPR, longer prosome length was also achievedby C. sinicus from May to July, while females in November developedshorter bodies but accumulated more energy for reproduction.     

BARNACLE-DWELLING EGOTYPES OF THREE BRITISH LITTORINA SPECIES AND THE STATUS OF LITTORINA NEGLECTA BEAN

On the basis of collections from SW Wales and W Ireland it isshown that small (<5 mm) black and/or white Littorina occurringin the barnacle zone on exposed shores, of the L. neglecta shelltype, comprise three species. From their reproductive anatomythese correspond to L. saxatilis, L. arcana and L. nigrolineata,and are interpreted as analogous barnacle-dwelling ecotypesof these species. This adds to the evidence that L. neglectais not a distinct species, but an ecotype of L. saxatilis. (Received 1 June 1992; accepted 23 June 1992)     

Egg production and hatching success in the peri-Antarctic copepod Calanus simillimus

In situ rates of egg production and hatching success are reportedfor Calanus simillimus, one of the most abundant calanoid speciesin peri-Antarctic regions, during the Italian ‘Italics’cruise in the Straits of Magellan in March-April 1995. Low fecundity(8.6 eggs female^–1 day^–1) and fecal pellet production(0.8 fecal pellets female^–1 day^–1) in this periodindicate that the species was feeding very poorly. Sixty-sixof the 126 females sampled did not produce eggs and 80 of thesedid not produce any fecal pellets during the 24 h period ofincubation. Striking abnormal naupliar and embryonic developmentwas recorded in 81.8% of the eggs spawned. Aberrant eggs didnot undergo normal cleavage, and failed to develop to hatching.Deformed nauplii were asymmetrical and presented strong anatomicalanomalies in the total body length and number of swimming appendages.These results are discussed in the light of recent findingson the causes of low hatching success in copepods.     

Field Measurements of the Gas Exchange of Woody Plant Species in Simulated Sunflecks

The photosynthetic responses of eight tree and shrub speciesto simulated sunflecks was measured in the field. The net carbonexchange (NCE) of Corylus avellana and Ulmus glabra increasedwith irradiance up to the maximum irradiance of 230 µmolm^–2 s^–1. The NCE of Fraxinux excelsior, Hedera helixand the sun and shade forms of Rhododendron ponticum saturatedat about 120 µmol m^–2 s^–1 whereas the NCEof Ilex aquifolium, Daphne laureola and Fagus sylvatica hadeffectively saturated at 27 µmol m^–2 s^–1. In all cases the quantum efficiency of NCE could be predictedfrom measurements of chlorophyll fluorescence and the maximumvalue for NCE from measurements of stomatal conductance. Therelationships were combined into a model for predicting NCE/irradiancecharacteristics. Corylus avellana L., Daphne laureola L., Fagus sylvatica L., Fraxinus excelsior L., Hedera helix L., Ilex aquifolium L., Prunus laurocerasus L., Rhododendron ponticum L., Ulmus glabra Huds., gas exchange, stomatal resistance, water use efficiency, chlorophyll fluorescence, quantum efficiency     

Food type induces different reproductive responses in the copepod Centropages typicus

Egg production and hatching success were determined for individualCentropages typicus fed two diatoms (Thalassiosira rotula andPhaeodactylum tricomutum) and two dinoflagellates (Prorocen-trumminimum and Gonyaulaxpolyedrd). Both reproductive responseswere strongly affected by food type. Females incubated withoutmales produced eggs with all diets, but fecundity was twiceas high with the larger T. rotula and G.polyedra cells. In contrast,hatching success was 2–3 times higher with the dinoflagellatediets. The presence of males did not enhance egg productionrates. Males also did not improve hatching success when thediet consisted of the diatom T.rotula. However, egg viabilitywas higher for couples fed the dinoflagellate G.polyedra, indicatingthat egg viability was possibly being controlled by both rematingand food type. Egg viability was artificially lowered by exposingnewly spawned eggs to high concentrations (10⁴–10⁹ pgC) of extracts from T.rotula, whereas the development of eggsproceeded normally at all concentrations of extracts from P.minimum.Blockage of egg development was not due to anoxia, but to thepresence of intracellular, deleterious chemical compounds indiatoms, suggesting that bottom-up prey control mechanisms underlieseasonal fluctuations in C.typicus populations at sea.     

The Mechanism of Action of Vernalization in Lathyrus odoratus L.

In the sweet pea (Lathyrus odoratus L.) genes Dn^l andDr^h controlthe production of a graft-transmissible substance which delaysflowering and promotes outgrowth of basal laterals. Seed vernalizationpromotes flowering and reduces lateral outgrowth in intact plantsand grafted scions of genotype DnⁱDn^l, suggesting that vernalizationreduces output of the Dnⁱ system, possibly by disrupting therelationship between chronological and plastochronic age. Whenlateral outgrowth and floral abortion are used as indicatorsof inhibitor levels, it can be shown that vernalized Dnⁱ plantspossess more inhibitor but initiate flower buds at a lower nodethan unvernalized dn plants. This supports the suggestion thatin regard to floral initiation vernalization also alters thesensitivity of the shoot apex to the flowering hormone(s). InLathyrus odoratus an hormonally based vernalization responseof considerable magnitude can be shown for day-neutral (dndn)lines, supporting the suggestion that vernalization also influencesthe level of a flower promotor. Lathyrus odoratus L., sweet pea, vernalization, flowering, branching, genotype, grafting     

A GENETIC ANALYSIS OF SYMPATRIC SUBPOPULATIONS OF THE SIBLING SPECIES LITTORINA SAXATILIS (OLIVI) AND LITTORINA ARCANA HANNAFORD ELLIS

A detailed analysis of shell characteristics (four parameters)and allozyme allele frequencies (eight to nine polymorphic loci)of sympatric Littorina saxatilis and Littorina arcana subpopulationsfrom Filey Brigg on the east coast of England supports theiridentification as closely related but separate species. Oneenzyme system, esterase-2, was nearly species diagnostic. Resultswere generally comparable with an earlier study of the two speciesfrom East Lothian, Scotland. Littorina saxatilis was again foundto be more variable than L. arcana. (Received 14 December 1983;     

SPAWNING SEASON AND EGG PRODUCTION IN FORTH ESTUARY RETUSA OBTUSA (MONTAGU) (GASTROPODA: OPISTHOBRANCHIA)

Retusa obtusa held individually in the laboratory carried largeoocytes in the gonad but did not spawn in December-late January.In late Jan-early Feb, however, the first few individuals producedbatches of 8–17 eggs and, by early February, most individualswere producing means of 2–4 eggs d^–1 and maximain late Feb of <7.9 d^–1. Spawning declined to low levelsin few individuals in late March. By early April, spawning hadvirtually ceased prior to the drastic loss of adults in thepopulation by May. Larger R. obtusa generally produced more eggs than smaller onesand the few specimens shorter than 3.8 mm produced markedlyfewer than those of 3.8–5.0 mm shell length. Twenty of the 34 R. obtusa held in the main breeding period(4 sets started 29 Jan-3 Mar) spawned in 3 or more successiveweeks whereas only 6 of them failed to spawn in 2 consecutiveweeks, so indicating that several successive batches of eggsare commonly produced. Yet in virtually all individuals (exceptat the start of breeding) more eggs were produced in the firstweek of captivity than subsequently. These first-week valuesimply total production by individuals in the spawning seasonof some 112-305 eggs comprising about 2.8-7.6 mg dry weightor, by the Forth estuary population, of 26880-73200 eggs m^–2comprising 0.67-1.83 g dry weight m^–2. (Received 19 August 1988; accepted 30 September 1988)