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)     

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)     

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 FINE STRUCTURE AND FUNCTION OF THE ANAL GLAND OF THE MURICID NUCELLA LAPILLUS (NEOGASTROPODA) (AND A COMPARISON WITH THAT OF THE TROCHID GIBBULA CINERARIA)

The fine structure of the anal gland of Nucella lapillus isdescribed, and compared with that of Gibbula cineraria. It isconcluded that the glands are not analogous in function andare unlikely to be homologous. The anal gland of Gibbula isa small rectal diverticu-lum near the anal papilla, lined bymucus-secreting cells. The evidence suggests that the largeanal gland of Nucella extracts macromolecules and cations fromthe blood, metabolizes or sequesters them in lyso-somes andultimately expels the resultant residual bodies by apocrinesecretion. It has large reserves of lipid and glycogen, andthe abundance of melanin indicates that it is a major site oftyrosine degradation. The invariable presence of bacteria inthe lumen, and of pits in the epithelium to house them, impliesa symbiotic relationship in which the bacteria metabolize anddegrade the cell debris, some of which is resorbed. The lossof the anal gland in bucci-nids and nassariids is accompaniedby great reduction in size and importance of the rectal sinus,and increase in complexity of the kidney. (Received 31 December 1991; accepted 23 January 1992)     

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;     

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)     

ALLOZYME SIMILARITY OF ATLANTIC AND PACIFIC SPECIES OF LITTORINA (GASTROPODA: LITTORINIDAE)

Four Littorina species from the Pacific (L. sitkana, L. brevicula,L. mandshurica, L. squalida) and four from the Atlantic (L.mariae, L. obtusata, L. saxatilis and L. littorea) were comparedelectrophoretically using 17 loci. Analysis of genetic identities showed that there were threegroups. The first group comprises L. sitkana, L. mariae, L.obtusata and L. saxatilis The second group consists of L. breviculaand L. mandshurica and the third one includes L. squalida andL. littorea. Our results support Reid's hypothesis of the originof Atlantic littorines from two Pacific ancestors. (Received 18 December 1991; accepted 30 March 1992)     

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)     

A PLANIMETRIC STUDY OF MORPHOLOGICAL VARIABILITY IN THE DIGESTIVE DIVERTICULA OF LITTORINA LITTOREA (LINNAEUS) AND MYTILUS EDULIS LINNAEUS

The inter-and intratubular morphological variability in thedigestive diverticula of Littorina littorea and Mytilus edulishas been investigated by planimetric procedures. Five parametershave been measured: mean epithelial thickness (MET), mean diverticularradius [MDR], mean luminal radius (MLR), MLR/MET ratio and MET/MDRratio. The results indicate that irrespective of the patternof tubule organization within the digestive diverticula (whethertubule types are clustered or not), variability between individualsis greater than that between zones of the digestive gland. Thishas implications for the design of sampling strategies in investigationsof the morphology of digestive diverticula in physiologicaland pathological studies. The variability in the epithelialthickness within diverticular sections is of minor relevancein assessing the overall condition of the digestive gland inthese studies, because variability in epithelial thickness betweentubules is significantly greater than within tubules in bothspecies. (Received 10 March 1989; accepted 16 October 1989)     

THE ANATOMY OF PACHYDERMIA LAEVIS (ARCHAEOGASTROPODA: 'PELTOSPIRIDAE')

The anatomy of Pachydermia laevis Warén & Bouchet,1989 is investigated. It is a deep-sea gastropod and restrictedto hydrothermal vents of the East Pacific Rise. Its anatomyresembles that of Melanodrymia aurantiaca Hickman, 1984 (Melanodrymiidae)in most respects, except that its gonopericardial duct opensinto the renopericardial duct, not into the peri-cardial chamber,and that it lacks a copulatory organ. Examination of M. aurantiacamain confirms earlier work. The two species have a cerebralbuccal connective fused with the cerebral ganglion (not freefrom it), a synapomorphy that has not been described for anyother archaeogastropod. The investigation suggests that Melanodrymiaand Pachydermia are closely related and together with Cyathermiidaeand Neomphalidae form a monophyletic group within the Neomphalina.No evidence was found to support earlier speculations aboutclose relations of Neomphalina to Viviparoidea. (Received 10 January 1996; accepted 3 April 1997)     

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;     

THE ANATOMY AND RELATIONSHIPS OF THE ORBITESTELLIDAE (GASTROPODA: HETEROBRANCHIA)

The anatomy of two genera of the marine gastropod family, Orbitestellidae,Orbitestella and Microdiscula, is described. They possess apigmented mantle gland and pallial renal organ, lack a ctenidiumand have a simple hermaphrodite genital system with an externalpenis. They lack buccal cartilages, have a simple oesophagus,a massive jaw and a crystalline style in the stomach. The Orbitestellidaeare shown to be primitive heterobranchs, perhaps related tothe northern-hemisphere freshwater family Valvatidae. (Received 9 September 1989; accepted 30 December 1989)     

THE ANATOMY AND RELATIONSHIPS OF A MARINE VALVATOIDEAN (GASTROPODA: HETEROBRANCHIA)

The anatomy of Cornirostra (new genus) pellucida (Laseron),a marine Indo-West Pacific gastropod, is described and is shownto be similar to that of the northern hemisphere, freshwaterValvatidae. There are, however, several differences betweenCornirostra and valvatids, notably in details of the anatomyof the reproductive system, the morphology of the head-foot,the spawn and in radular characters that warrant family-levelseparation for Cornirostra. Tomura bicaudata (Pilsbry and McGinty) from Florida is alsoa member of the family. (Received 9 September 1989; accepted 30 December 1989)     

ALLOZYME VARIATION AMONG POPULATIONS OF BIOMPHALARIA CAMERUNENSIS (BOETTGER, 1941) (GASTROPODA: PLANORBIDAE) IN CAMEROON

Allozyme electrophoresis was used to study genetic diversityamong populations of the freshwater, hermaphroditic snail Biomphalariacamerunensis in Cameroon. Five of 19 loci studied in 15 enzymesystems were polymorphic. Intrapopulation variation was evidentin 8 of 12 populations sampled and heterozygotes were presentin 6 of these. Two populations with large sampled sets werepanmictic. Although results for the remaining 4 populationswith heterozygotes present were inconclusive, our findings supportthe hypothesis that Biomphalaria are out-breeders. The allelesfor two loci, asparate aminotransferase-1 (AAT-1) and 6-phosphogluconatedehydrogenase (6-PGD), showed distribution patterns that suggestthey may be limited by climate or habitat. The slow migratingAAT-1 was the only allele found in the mountainous regions ofthe southwest. This region falls under the Cameroon-type climateand is the wettest region in the country. The faster allelewas restricted to the lowland rain forest of the south. For6-PGD the fast allele was found throughout the range of B. camerunensis,but the slower allele was found only in the mountainous regionswithin the Cameroon-type climate. (Received 27 April 1989; accepted 9 September 1989)     

THE FINE STRUCTURE AND FUNCTION OF THE FOOT OF NASSARIUS KRAUSSIANUS, A GASTROPOD MOVING BY CILIARY LOCOMOTION

Light and electron microscope studies of the oval foot of Nassariuskraussianus (Dunker) showed that the columellar muscle is asingle extrinsic muscle and dorsoventral, transverse and longitudinalfibres are the principal intrinsic muscles. The fine structureof the columellar muscle and the intrinsic musculature is describedand shown to be similar to that of other gastropods. The presenceof abundant mitochondria in a snail moving by cilia may be relatedto the rapid and powerful movements of the foot made in escapingpredators. All pedal muscle consists of smooth muscle fibresand collagen was observed to be particularly associated withthe intrinsic fibres. The foot maintains a constant area ofpedal sole during ciliary locomotion by means of antagonismbetween the intrinsic muscle fibres. The possible role of collagenin this process is discussed. (Received 6 March 1989; accepted 15 June 1989)     

THE PHYSIOLOGY AND FEEDING BEHAVIOUR OF TWO MARINE SCAVENGING GASTROPODS IN HONG KONG: THE SUBTIDAL BABYLONIA LUTOSA (LAMARCK) AND THE INTERTIDAL NASSARIUS FESTIVUS (POWYS)

Aspects of the physiology and feeding behaviour of the scavengingsnails, Babylonia lutosa and Nassarius festivus have been studiedand compared. N. festivus has a wider physiological tolerancethan B. lutosa; upper lethal temperatures being 31.5°C and28.5°C respectively and lower lethal salinities 15.5 and11.5. N. festivus was also more resistant to desiccation. N.festivus is adapted to an intertidal environment that is morevariable than the subtidal habitat of B. lutosa. The metabolicrate of adult N. festivus was two to three times higher thanthat of adult B. lutosa, the specific oxygen consumption rateof the former being 7.91±0.49x10^–6 mol of O₂.g^–1wet wt.hr^–1. This correlates with estimates of consumptionobtained for the two animals, B. lutosa eating, on average,6.57% of its wet tissue weight per day, N. festivus 15.33%.Standardization of the data for 2 g animals of both species,however, reveals no significant difference in terms of oxygenconsumption and thus the enhanced consumption by N. festivussimply reflects its smaller size. Both species are opportunisticfeeders and can survive long periods of starvation (>100days). The feeding strategies of the two are also different,N. festivus eating faster than B. lutosa, possibly because ofthe more ephemeral nature of carrion in the intertidal. Starvationaffected the feeding behaviour of N. festivus more than B. lutosaand N. festivus was able to perceive carrion from greater relativedistances than B. lutosa. (Received 27 February 1989; accepted 24 July 1989)     

A NEW SPECIES OF OXYCHILUS FROM THE LIGURIAN APENNINES (ITALY) (PULMONATA: ZONITIDAE)

A new species of Oxychilus Fitzinger from the Ligurian Apennines(Italy) is described. The new species is particularly characterizedby its penial complex which has a very slender proximal peniswith variable inner sculpture and a widened distal penis. Theproximal penis inner structure consists of a series of longitudinalpleats which may often be crenulated, undulated or fragmentedto form rows of papillae, providing new evidence to supportthe opinion that this is not a valid diagnostic character forseparating Oxychilus into the nominal subgenus and OrtiziusForcart. Consequently Ortizius is herein proposed as a juniorsynonym of Oxychilus s. str. (Received 5 July 1990; accepted 4 March 1991)     

STRUCTURE OF THE CEPHALIC TENTACLES OF SOME SPECIES OF PROSOBRANCH LIMPET (PATELLIDAE AND FISSURELLIDAE)

The morphology and ultrastructure of the cephalic tentaclesand eye (optic vesicle) of some patellid and a fissurellid limpetspecies are described. The epithelium of the tentacle bearsciliated cells which have neural connections suggesting a sensoryfunction. In patellid limpets the density of these cells variesbetween species, with the greatest density (18 ciliated tufts.100 µm^–2) being recorded in the territorial limpet,Patella cochlear. The surface of the tentacle of a fissurellidlimpet, Fissurella mutabilis, is papillate, which contrastswith the smooth surface of a patellid tentacle. Ciliated tuftsare borne at the tip of most papillae. Movement of the tentacleis controlled by longitudinal muscle and a radial muscle-collagentissue association, which function as a muscular hydrostat.The eye of a patellid consists of a simple open vesicle anda retina which is comprised of one type of cell. By contrastthe optic vesicle of a fissurellid contains a lens, enclosedby a cornea and a retina which is composed of two types of cell,pigmented and photoreceptive. The ultrastructural features ofthe cells resemble those described for other molluscan eyes. (Received 5 June 1989; accepted 16 August 1989)     

ASYNCHRONOUS REPRODUCTIVE ACTIVITY IN THE BROADCAST SPAWNER CELLANA CAPENSIS (GMELIN, 1791) (GASTROPODA: PATELLIDAE)

Short-term fluctuations in the reproductive condition of thelimpet Cellana capensis are described. Gravimetric and histologicalanalyses of gonadal development were carried out on samplescollected at 4 day intervals over a 3 month period from theMkambati Nature Reserve, Transkei. The results indicate a distinctlack of reproductive synchrony within the population, whichapplied not only to gametogenic development per se but alsoto spawning activity. Some of the spawning events identifiedduring the study period were more marked than others, thesemay reflect a greater degree of synchronization in activity.Results presented suggest that C. capensis is probably a ‘partial’rather than a ‘complete’ spawner. The implicationsof asynchronous spawning activity for the reproductive successof broadcast spawners are also discussed. (Received 6 December 1988; accepted 15 February 1989)     

Microsatellite cross-species amplification in the genus Littorina and detection of null alleles in Littorina saxatilis

Microsatellite DNA is widely used as population genetic marker,but the cost of using microsatellites is high, as they usuallyneed to be developed and optimized for each species separately.Cross-species amplification of microsatellites is thereforecommonly applied to bring down the cost, but it can also involvegenotyping errors. We studied cross-species amplification ofmicrosatellites in four species of the Atlantic group of Littorina(Neritrema): L. saxatilis (Olivi, 1792), L. obtusata (Linnaeus,1758), L. fabalis (Turton, 1825) and L. arcana Hannaford Ellis,1978 to investigate whether markers originally developed fora more distantly related Pacific species [L. subrotundata (Carpenter,1864)] suffered from more amplification problems than markersdeveloped for one of the species in the Atlantic group (L. saxatilis).We also compared variation in amplification success among thespecies and among different regions in the NE Atlantic. Approximatelyhalf of the 12 primers developed for L. subrotundata and theseven primers developed for L. saxatilis were successfully amplifiedin other species of the subgenus. The success was dependenton phylogenetic distance among species within the subgenus.On the other hand, the variation in performance of the locibetween geographically remote populations of the same specieswas as high as variation among the species. In earlier studiesstatistical analyses indicated that several loci showed a heterozygotedeficiency due to null alleles. The presence of null alleleswas confirmed by a segregation analysis of the microsatelliteloci in eight half-sib families of L. saxatilis. (Received 2 April 2007; accepted 19 November 2007)