TENACITY OF ATTACHMENT IN TWO SPECIES OF LITTORINID, LITTORINA LITTOREA (L.) AND LITTORINA OBTUSATA (L.)

Here we present data concerning the ability of two littorinidsto remain attached to a slate substratum as a dislodging forceis applied vertically. Specimens of Littorina littorea (L.)and L. obtusata (L.) were attached to a simple pulley and sandflowsystem which applied force to the animals at a constantly increasingrate until the animals released from the substratum. Three aspectsof tenacity were measured: force required to remove the winklesfrom the substratum; time taken for removal; and final displacementat removal. For L. littorea the mean force required (1.09 N),the mean displacement (8.9 mm) and the mean time taken (185s) were all found to be significantly greater than those recordedfor L. obtusata (means = 0.60 N, 7.5 mm and 102 s, respectively).However, when animal size was taken into account there was nosignificant difference in either force or time required forremoval, although L. obtusata showed a greater displacementthan did L. littorea. Animal dry flesh weight was not a usefulpredictor of any of the tenacity variables. Rugosity of theslate substrata was shown to have a significant effect on alltenacity variables, greater forces being required to removeboth species of winkle from smooth substrata than from roughsubstrata. This also generally held true when animal size wasaccounted for. The results of these experiments, although insufficientto identify the mechanism(s) of attachment, do discount themechanism of ‘muscular grip’, since greater tenacitywould be expected on rough substrata. Whether winkles were immersedor emersed had no significant effect on any of the tenacityvariables. Recorded tenacities suggest that many L. littoreaand L. obtusata would be dislodged by the direct forces createdby breaking waves. (Received 5 August 1996; accepted 18 November 1996)     

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)     

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)     

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;     

VARIATION IN THE OCCURRENCE OF ABNORMAL EMBRYOS IN FEMALES OF THE INTERTIDAL GASTROPOD LITTORINA SAXATILIS OLIVI

The incidence of abnormally developed embryos within the brood-pouchof female L. saxatilis was very variable, ranging from 0–100%of embryos being abnormal. A minor part of the total variationwas attributed to between-shore variation, while 82% reflectedwithin sample differences between females. Possible factorsresponsible for the variation in embryo abnormality shown bythis and other investigations, are discussed and it is concludedthat genetic rather than non-genetic factors are important. (Received 8 November 1983;     

MORPHOLOGICAL DIVERGENCE AND EVIDENCE FOR REPRODUCTIVE ISOLATION IN LITTORINA SAXATILIS(OLIVI) IN NORTHEAST ENGLAND

Current work at Old Peak (Ravenscar) on the northeast coastof England indicates that boulder-dwelling Lutonna saxatilis(Olvi) comprise at least two distinct forms, inhabiting high-and mid-shore boulders Using multivariate morphometrics we showthat these forms are significantly different in shape. Rareintermediates between these two distinct forms were occasionallyfound, usually with the low shore animals The two forms showdifferent reproductive strategies. High shore females (L. saxatilisH) mature at a smaller size and produce a smaller number oflarge eggs giving rise to large crawlaways The mid-shore form(L. saxatilis M) matures at a larger size and produces a largenumber of small eggs which give rise to small crawlaways. Anovel observation is that intermediate females (L. saxatilisI) contain eggs of both sizes, giving rise to a bimodal distributionof egg sizes within the brood pouch. The usual incidence ofabortion (observed in H and M forms) is Very low (<1%) However,in the intermediates the large eggs showed a high incidenceof abortion (about 60%) and the incidence of abortion in thesmall eggs was also increased (about 30%) Thus there appearsto be a partial reproductive barrier between the two forms,presumably resulting in reduced fitness due in the intermediatesto reproductive dysfunction. These findings extend to othershores in the region, over total distance of about 66 km. Thefacts are suggestive of a hitherto unrecognised reproductivebarrier with L. saxatilis, the nature and strength of whichis under investigation ^*Present address: University College Scarborough. Filey Road.Scarborough Yo11 3AZ. (Received 12 June 1995; accepted 15 September 1995)     

FUNCTIONAL MORPHOLOGY OF THE RECTUM IN THE MARINE GASTROPOD LITTORINA SAXATILIS (OLIVI) (PROSOBRANCHIA:LITTORINOIDEA)

The ultrastructure of the rectum of the marine prosobranch gastropod,Littorina saxatilis suggests that it may have an additionalrole to the voiding of faecal pellets. It consists of two epitheliallayers, separated by a blood space. The inner epithelial layeris greatly folded and consists of metabolically highly activecells containing a wide variety of inclusions, and showing evidenceof secretory activity at their luminal surfaces. The outer epitheliumcontains a greater diversity of cell types, also with a largenumber of different inclusions present. Unlike some other prosobranchgastropods, a distinct rectal gland is absent in L. saxatilis.The entire rectal epithelium is glandular, and it appears thatthe rectal epithelial cells produce a number of secretions.This may indicate a role in absorption, the removal of materialfrom the blood, or the addition of a final covering to the faecalpellets. (Received 6 February 1989; accepted 6 March 1989)     

ANNUAL AND LUNAR REPRODUCTIVE CYCLES IN LITTORINA SAXATILIS (OLIVI) AND DIFFERENCES BETWEEN BREEDING IN THE MARINE FIRTH OF FORTH AND THE FORTH ESTUARY

Gonads were most commonly reduced in October and fullest inMar-Jun with more pronounced seasonal differences in the estuarythan in the firth. Brood-pouches accordingly were fullest inspring-early summer and emptiest in autumn-early winter. Eggsand early embryos predominated in brood-pouches of early springbut were few in Aug-Oct whereas well-developed juveniles werefew in early spring, increased in spring-summer and came topredominate in Jun-Nov. Brood-pouch loads varied with size offemale but, for a standard 10 mm female, were significantlylower in the 2 estuarine sites than in the firth. Changes inmean brood-pouch count varied only 2.5-fold during the yearin the firth but 4.10-fold in the estuary. In vitro development of eggs to shelled juveniles with the velumresorbed took some 68 d at 10°C and 32%. Eggs and late brood-pouchjuveniles were significantly heavier in the estuary than inthe firth (dry weight, ash-free dry weight (AFDW), ash weight).AFDW changed little during development while ash weight (mostlyshell) increased 3.5-4.4-fold. In a laboratory tide tank, femalesreleased 60 d means of <1.3 juveniles d^–1; more inmid- and late year than in Jan-Mar, more from upshore firthfemales than from low shore or estuary females, more when heldat high tide-tank levels than at lower levels, and generallyat rates comparable to release in containers held at the siteof origin. Egg production in laboratory females did not keeppace with release of young and, at the end of trials, brood-pouchcounts were mostly lower than in contemporary free winkles.Females that had released most (e.g. at high tank levels) containedleast brood-pouch embryos and young subsequently. Release of young in the tide tank was markedly faster at timesnear new moon (often reaching or exceeding 2 female^–1d^–1) than near full moon (often <1 female^–1 d^–1).This lunar fluctuation was clearest in fast-releasing sets heldat high- or mid-tide tank levels. It was not observed in Jan-Mar(when release from most females was slow) nor in females atlow tank levels. (Received 3 September 1990; accepted 8 November 1990)     

ON THE OCCURRENCE OF THE PROSOBRANCHS LITTORINA NERITOIDES, L. MARIAE AND L. OBTUSATA IN DENMARK

Littorina neritoides occurs on piers and large stones on thebeaches in northern Jutland, but may not be able to toleratecold winters. L. obtusata and L. mariae are common on Fucus,but only L. mariae penetrates deeply into the fjords and sounds. (Received 10 December 1979;     

PHENOTYPIC DIFFERENTIATION IN LITTORINA SAXATILIS OLIVI (MOLLUSCA, PROSOBRANCHIA) IN A SMALL AREA ON THE SWEDISH WEST COAST

The Littorina saxatilis species-complex exhibits two distinctmorphs in the area investigated. Differences in shell morphologyand number of penial glands are statistically significant (P<0.02)between populations from exposed and sheltered habitats. However,in intermediate environments intermediate specimens as wellas typical specimens of both morphs are found. The differencesare considered intraspecific and at least partly geneticallydetermined, and possible selection pressures responsible arediscussed. (Received 8 February 1981;     

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)     

INCIPIENT REPRODUCTIVE ISOLATION BETWEEN TWO SYMPATRIC MORPHS OF THE INTERTIDAL SNAIL LITTORINA SAXATILIS

The study of speciation in recent populations is essentially a study of the evolution of reproductive isolation mechanisms between sub-groups of a species. Prezygotic isolation can be of central importance to models of speciation, either being a consequence of reinforcement of assortative mating in hybrid zones, or a pleiotropic effect of morphological or behavioral adaptation to different environments. To suggest speciation by reinforcement between incipient species one must at least know that gene flow occurs, or have recently occurred, and that assortative mating has been established in the hybrid zone. In Galician populations of the marine snail Littorina saxatilis, two main morphs appear on the same shores, one on the upper-shore barnacle belt and the other in the lower-shore mussel belt. The two morphs overlap in distribution in the midshore where hybrids are found together with pure forms. Allozyme variation indicates that the two parental morphs share a common gene pool, although within shores, gene flow between morphs is less than gene flow within morphs. In this study, we observed mating behavior in the field, and we found that mating was not random in midshore sites, with a deficiency of heterotypic pairs. Habitat selection, assortative mating, and possibly sexual selection among females contributed to the partial reproductive isolation between the pure morphs. Sizes of mates were often positively correlated, in particular, in the upper shore, indicating size-assortative mating too. However, this seemed to be a consequence of nonrandom microdistributions of snails of different sizes. Because we also argue that the hybrid zone is of primary rather than secondary origin, this seems to be an example of sympatric reproductive isolation, either established by means of reinforcement or as a by-product to divergent selection acting on other characters.     

THE EGG MASS OF LITTORINA OBTUSATA AND LACUNA PALLIDULA (GASTROPODA : PROSOBRANCHIA)

The egg masses of Littorina obtusata and of Lacuna pallidulahave been compared with regard to size, shape, egg number, eggsize and substratum. The structure and histochemistry of thespawn of the two species has been investigated and observationson the laying process in L. obtusata are also recorded. ^* Present address: Portway School, Penpole Lane, Shirehampion,Bristol, BS11 0EB. (Received 12 October 1977;     

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;     

THE RHYTHMIC SPAWNING BEHAVIOUR OF LITTORINA LITTOREA (L.)

Daily intertidal plankton samples have established a semi-lunarperiodicity of spawning in Littorina littorea (L.). Within thegeneral pattern of seasonal spawning, the density of egg capsulesin the surface waters and the state of development of the embryosshow that oviposition takes place on both full and new moonspring tides (Received 25 August 1978;     

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)     

MORPHOLOGICAL DIFFERENTIATION AND GENETIC COHESIVENESS OVER A MICROENVIRONMENTAL GRADIENT IN THE MARINE SNAIL LITTORINA SAXATILIS

The marine gastropod Littorina saxatilis has different ecotypes in shores only a few meters apart. This has both taxonomic and evolutionary implications. Here we report on an extreme type of within-shore dimorphism in shell characters. In the wave-exposed rocky shores in northwestern Spain, we found one form of L. saxatilis in the upper-level barnacle zone. It had a white, ridged shell, with black bands in the grooves. Another form confined to the lower-shore mussel belt had a smooth shell that was either white and tessellated or darkly colored. These two forms cooccured in a narrow midshore zone together with individuals that had combined characters, but were present in low frequencies (11%–29%). We used principal-component analysis of metric shell characters to study variation in shell size and shape. We found that the upper-shore form was larger than the lower-shore form. We also found small but significant differences in shell shape. Experiments in a common laboratory environment suggested the differences in shell ornamentation and color are inherited, but the individuals did not develop the morph-specific characters until a shell height of about 3 mm. The occurrence of mainly two distinct forms may suggest the presence of two species that hybridize. An analysis of five polymorphic enzyme loci in populations of snails from three geographically separated sites indicated, however, that there was no positive correlation between morphological distances and genetic distances among populations on a geographic scale (tens of kilometers). Thus, we rejected the hypothesis of two species. However, on a microgeographic scale (meters), genetic differentiation between groups with the same form was less than differentiation between forms. This indicated a partial barrier to gene flow between the two forms, and preliminary mate choice data suggested this was caused by nonrandom mating in the midshore zone of overlap.     

FACTORS CONTRIBUTING TO SPATIAL HETEROGENEITY IN THE ABUNDANCE OF THE COMMON PERIWINKLE LITTORINA LITTOREA (L.)

Although Littorina littorea (L.) exhibits a relatively consistentpattern of vertical distribution throughout the North Atlantic,ranging from the mid-intertidal to the shallow subtidal zone,its horizontal distribution and abundance are highly variable.In this study, we first described the snail's horizontal distributionpatterns on Appledore Island, ME, USA and then asked whetherwave exposure, rugosity, predator density (e.g. Carcinus maenasand Cancer borealis), the percentage of the substrate coveredby Ascophyllum nodosum, Chondrus crispus, Polysiphonia spp.,and ephemeral green algae, or the percentage of uncovered substrate(bare rock) were correlated with L. littorea abundance in theintertidal zone (0.6 to 0.0 m relative to Mean Lower LowWater [MLLW]) and the shallow subtidal zone (–1.5 to –3.0 mMLLW) at nine study sites. Intertidal densities of L. littoreawere highly variable across sites, ranging from 0 to >600 m^–2.In this zone, L. littorea density showed a significant positivecorrelation with rugosity and percentage of bare rock. Densitieswere very low in the subtidal zone (range: 0–19 m^–2)and varied little among sites. Exploratory multiple regressionanalysis suggested that L. littorea density was positively correlatedwith the density of C. maenas in the shallow subtidal zone.Additionally, snails in the subtidal zone had thicker shellsthan snails of the same size in the intertidal zone. Our resultssuggest that structural elements of the habitat, such as rugosityand percentage of uncovered substrate, are among the most importantpredictors of L. littorea abundance on moderately protected,rocky intertidal shores. (Received 9 February 2005; accepted 10 August 2005)     

THE CHROMOSOME NUMBER AND POSSIBLE ANCESTRY OF PELLAEA WRIGHTIANA

Knobloch , Irving W. (Michigan State U., E. Lansing), and D. M. Britton . The chromosome number and possible ancestry of Pellaea wrightiana . Amer. Jour. Bot. 50(1): 52–55. Illus. 1963.—Meiotic and somatic chromosome number determinations show that Pellaea wrightiana Hook, is a triploid (2n = 87) of hybrid origin. P. ternifolia (Cav.) Link (4x) and P. longimucronata Hook. (2x) (2n = 58) are considered to be the probable parents.