ALLOZYME COMPARISON OF LITTORINA SPECIES FROM THE NORTHWESTERN PACIFIC

Three unknown Littorina species were revealed by allozyme studyof snails from the northwestern Pacific. One of them was foundin Kasatka Bay (Pacific coast of Iturup Island, south KurileIslands) together with L. sitkana, which it resembled in anatomicalfeatures, but the genetic distance between these species wasD=0.622. The second one was found in Tauiskaya Inlet (northernpart of Okhotsk Sea) and was again similar to L. sitkana inmorphological and anatomical features, but D=1.160. The thirdspecies was found in Egvekinot Inlet (Anadyrskiy Gulf, BeringSea) in sympatry with L. natica, to which was close genetically(D=0.256) although it differed in shell shape. (Received 4 April 2005; accepted 26 August 2005)     

RADULAR PRODUCTION RATES IN TWO SPECIES OF LACUNA TURTON (GASTROPODA: LITTORINIDAE)

The molluscan radula is a dynamic organ, both in terms of itsuse and production. New rows of teeth are constantly producedat the posterior end of the radula, while older, worn teethare shed anteriorly, producing a dynamic equilibrium. We useda cold-shock to mark the radular ribbon and measure tooth rowproduction rates in two gastropod species, Lacuna vincta (Montagu)and L. vanegata Carpenter. We found that the average tooth rowproduction rate at 10–11°C did not differ betweenthese two species, and was 2.94 (SE = 0.002) rows per day forLacuna vincta and 2.97 (SE = 0 002) for L. vanegata Inter-individualvariability in production rate was very low, and was correlatedwith shell length, smaller individuals had slightly higher productionrates. The total length of the radular nbbon varied greatlyamong individuals, ranging from 47 to 94 (2.57 to 5.68 mm) rowsin L vincta and 53 to 99 rows (2.80 to 7.14 mm) in L vanegata,and was only somewhat correlated with the length of the shelLThis great variability will result in large differences amongindividuals in the time it takes to replace the radula totally,from 14.96 to 35.44 days in L vincta and from 17 43 to 39 69days in L. vanegata. (Received 1 September 1995; accepted 20 November 1995)     

A NEW OVOVIVIPAROUS SPECIES OF TECTARIUS (GASTROPODA: LITTORINIDAE) FROM NIUE, SOUTH PACIFIC, WITH A MOLECULAR PHYLOGENY OF THE GENUS

A new ovoviviparous littorinid gastropod, Tectarius (Echininiopsis)niuensis, from Niue, west of the Cook Islands, is described.This is distinguished from the only other ovoviviparous memberof the genus, T. (E.) viviparus (Rosewater, 1982) from the MarianaIslands, here redescribed in detail. The new species is remarkabletor its high-level habitat in the littoral fringe on wave-exposedkarstic limestone cliffs, for its variation in shell shape accordingto tidal level, shell colour polymorphism, calcified operculumand penis with a single mamilliform penial gland. In a cladisticanalysis of morphological characters, including single representativesof each of the three other subgenera of Tectarius (Tectarius,Echininus, Tectininus), these two ovoviviparous species appearas sister-taxa. This is confirmed by a molecular phylogeneticanalysis of the same species, based on the sequence of a portionof the 16S ribosomal RNA mitochondrial gene. Neither analysisunequivocally confirms the monophyly of Tectarius. The divergenceof DNA sequences within Tectarius suggests that the genus arosein the Upper Cretaceous, much earlier than the oldest (UpperEocene) fossils. Only 4 of the 175 species of Littorinidae areknown to be ovoviviparous (with brooding through metamorphosis)and the possible adaptive significance of this type of developmentis discussed. Hitherto, its rarity had been explained by earlyextinction of poorly-dispersed brooding taxa. However, ovoviviparitymay have persisted in Echininiopsis for at least 35 millionyears, and has not precluded colonization of islands 6300kmapart. (Received 17 September 1996; accepted 15 November 1996)     

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)     

ALLOZYME COMPARISONS AMONG SPECIES OF THE BULINUS FORSKALII GROUP (GASTROPODA: PLANORBIDAE) IN CAMEROON

Allozyme comparisons were made for adult Bulinus forskalii,B. camerunensis, and B. senegalensis from populations in Cameroonusing starch gel electrophoresis. Fifteen loci were examinedbut no differences were found between B. forskalii and B. camerunensis,bringing into question the validity of B. camerunensis as aseparate species. Bulinus senegalensis differed from B. forskaliiand B. camerunensis in allozymes for acid phosphatase, alpha-glycero-phosphatedehydxogenase, hydroxybutyrate dehy-drogenase, and phosphoglucomutase.No differences were found, as reported elsewhere, between B.forskalii and B. senegalensis for aspartate amino-transferase,isocitrate debydrogenase, phosphogluco-isomerase, and xanthineoxidise. No polymorphism was seen in B. camerunensis or variousB. senegalensis populations examined. Three alleles for isocitratedehydrogenase were observed for B. forskalii. Mixed populationsof B. forskalii and B. senegalensis were found at four sitesbut no evidence of hybridization between these species was found. (Received 9 May 1988; accepted 30 August 1988)     

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;     

ALLOZYME COMPARISONS AND PLOIDY LEVELS AMONG SPECIES OF THE BULINUS TRUNCATUS/TROPICUS COMPLEX (GASTROPODA: PLANORBIDAE) IN CAMEROON

Allozyme electrophoresis, ploidy level, and, to a lesser extent,anatomy and shell morphology were used to characterize snailsof the Bulinus truncatus/tropicus complex in Cameroon. Starchgel electrophoresis revealed no intrapopulation variation within20 populations examined for 14 allozyme loci in 12 enzyme systems.Interpopulation differences in allozyme patterns were used todivide the populations studied into five groups. Each groupis distinguished by a unique combination of zymograms for hydroxybutyratedehydrogenase and isocitrate dehydrogenase. Four of these groupsare tetraploids and are identified as B. truncatus. Populationsof the most commonly found B. truncatus group are generallyaphallic, are frequently associated with transmission of S.haematobium, and generally occur at altitudes below 800 m abovesea-level. Populations of B. truncatus belonging to the otherthree groups are usually euphallic, not associated with S. haematobiumtransmision, and generally occur at altitudes above 1000 m.These four groups may represent distinct biotypes of B. truncatus.Populations belonging to the fifth group are diploid, euphallic,not associated with S. haematobium transmission, and occur inhabitats above 1000 m. This is the first confirmed report ofdiploid B. truncatus/tropicus-complex snails from this regionof Africa. They are tentatively identified as B. natalensis,a species previously thought to be limited to East and SouthernAfrica. (Received 31 October 1988; accepted 20 February 1989)     

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;     

POPULATION BIOLOGY OF THE TRANS-ARCTIC EXCHANGE: MtDNA SEQUENCE SIMILARITY BETWEEN PACIFIC AND ATLANTIC SEA URCHINS

MtDNAs from 2 protein coding regions comprising 576 base pairs were sequenced from 17 individual sea urchins of the species Strongylocentrotus pallidus collected from the north Pacific and north Atlantic oceans. Twelve of 17 individual sequences were identical. Two of these were further sequenced in a third, 441 base pair region, and were also found to be identical. We show how to interpret these results using a simplified Markov model of mtDNA evolution at silent sites. The model was calibrated using 3 urchin species with a published fossil record, and shows that identical S. pallidus mtDNAs in different oceans shared a common ancestor at most 90,000–150,000 years ago (95% confidence interval of upper limit of divergence). The Markov model, used to examine patterns of genetic distance within and between species, shows unexpected variation in the rate of base substitution. The rate of change of G's at fourfold sites is nearly 20 times greater than the rate of change of C's. At twofold sites, this range is less extreme, although purines consistently have a higher rate of change than pyrimidines. Striking genetic similarity and recent genetic exchange between oceans for these urchins is in marked contrast to most other trans-Arctic marine populations, which usually show morphological and genetic differentiation at the species or subspecies level. Recent fossil evidence shows that the north Atlantic and northeastern Pacific have been the scene of radical faunal changes during the Pliocene and Pleistocene. The genetic results presented here extend this conclusion to intraspecific patterns of genetic variability, and direct attention to the northwest Pacific where higher productivity and less environmental change may have left a heritage of greater marine genetic diversity.     

DISTRIBUTION AND ABUNDANCE OF GYMNOSOMATA (GASTROPODA: OPISTHOBRANCHIA) IN THE SOUTHWEST ATLANTIC

The distribution and abundance of gymnosome gastropods in theArgentine Sea and Brazil—Malvinas Confluence during 1978–1979and 1988 were studied. The collections analyzed included 768quantitative samples obtained between 48°W and the coast,and from 35°S to 55°S. Two species were found. Spongiobranchaeaaustralis was the most frequent and abundant (up to 730 per1000 m3); its presence in the area was associated with the coreof the Malvinas Current. Clione antarctica was less abundant(maximum abundance: 230 per 1000 m³) and was also associatedwith the Malvinas Current. The geographic ranges of both speciesin the area are wider than previously described. Since the rangeof S. australis in the area extends far from the range of itsprey Clio, it is not clear whether S. australis can feed onthe thecosomatous pteropod Limacina (and not only on Clio, asdescribed in the bibliography) or it starves in that area. Duringthe 1978–1979 annual cycle, the abundance of both speciesfollowed neither the abundance patterns of their prey nor ofthe total zooplankton, and differed from each other. The residencetime of swarms of both gymnosomes were shorter than one month.As a general pattern, the aggregates are rapidly transportednorthward by the Malvinas Current and also penetrate the outershelf water, but they remain there only during a short periodand cannot preclude the final expatriation. So, the abundanceof gymnosomes in the area depends on passive migration morethan intrinsic population factors. (Received 9 July 1997; accepted 15 December 1997)     

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;     

TRANSFER EXPERIMENT SUGGESTS ENVIRONMENTAL EFFECTS ON THE RADULA OF LITTORARIA FLAVA (GASTROPODA: LITTORINIDAE)

Phenotypic variation in radulae has been studied in severallittorinid species because of this organ's intrinsic relationshipwith diet and, consequently, with the environment. In this work,we compared the radulae of the Brazilian species Littorariaflava found in mangroves and on rocky shores. Individuals ofL. flava showed marked differences in the shape of the cuspsamong samples from rocky shore and mangrove. In a transfer experiment,the shape of the radula changed within 40 days. A differentresponse was observed in individuals transferred to mangrove,where two different phenotypes were found, suggesting eitherintrapopulational variation in the responses to change of environmentalconditions, or that some snails showed a slow reaction to theenvironmental changes. The alterations could be attributed toecophenotypic plasticity. Analysis of variance showed that thelength of the radula in L. flava was strongly influenced bythe substrate (F_6,22=17.13, P<0.000), but apparently notby the transfer experiment. (Received 3 May 2005; accepted 18 July 2005)     

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)     

EVOLUTION OF ATLANTIC AND PACIFIC COD: LOSS OF GENETIC VARIATION AND GENE EXPRESSION IN PACIFIC COD

An allozyme investigation of 41 protein-coding loci in two morphologically similar fishes, Atlantic and Pacific cod, indicates that Pacific cod experienced a severe population bottleneck that led to the loss of gene diversity and gene expression. Pacific cod possesses a significantly lesser amount of gene diversity (H = 0.032) than Atlantic cod (H = 0.125) and lacks gene expression for Me-3. Allele-frequency distributions differ between species as predicted by neutral theory: Atlantic cod has a U-shaped distribution, which is expected for populations in drift-mutation equilibrium, whereas Pacific cod has a J-shaped distribution with an excess of low-frequency alleles. This excess may be explained by the appearance of new alleles through mutation which have not yet reached intermediate frequencies through drift. The population bottleneck in Pacific cod was most likely associated with founder populations that dispersed into the Pacific Ocean after the Bering Strait opened. Under the molecular-clock hypothesis a Nei genetic distance of 0.415 (based on 41 loci) suggests that Pacific cod dispersed into the Pacific Ocean soon after the Bering Strait opened in the mid-Pliocene, 3.0 to 3.5 million years ago.     

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)     

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)     

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)     

ALLOZYME VARIATION AMONG POPULATIONS OF BIOMPHALARIA PFEIFFERI (KRAUSS, 1848) (GASTROPODA: PLANORBIDAE) IN CAMEROON

Allozyme electrophoresis was used to study genetic diversityamong the freshwater, hermaphroditic snail Biomphalaria pfeifferiin Cameroon. Four of 19 loci studied were polymorphic. The allozymesfor two loci, aspartate aminotransferase-1 (AAT-1) and isocitratedehydrogenase (IDH), showed distribution patterns related toclimatic zones. AAT-1¹⁰⁰ and IDH¹⁰⁰ occurred predominantly inthe Tropical Climatic Zone, located in the northern half ofthe country. AAT-1¹⁴⁰ and IDH⁹⁰ were common in the EquatorialZone. Intrapopulation variations occurred in 5 of 19 populationsampled. Only one population, polymorphic for AAT-1 locus, waspanmictic. The other 4 populations were not in Hardy-Weinbergequilibrium. The deficiency of heterozygotes in those populationsis unusual for a member of the genus Biomphalaria and may bedue to a predominance of self-fertilization. Other explanationsinclude ecological factors or the presence of segregated sub-populationsthat do not outcross. This latter hypothesis is supported bythe total absence of heterozygotes for PGM alleles. (Received 24 August 1989; accepted 14 September 1989)