Multiple paternity in Littorina obtusata (Gastropoda, Littorinidae) revealed by microsatellite analyses.

Parental identity for juvenile Littorina obtusata was determined from three egg masses by means of microsatellite DNA markers. Results confirm that the attendant adult female in each case was the dam of the offspring and that at least 4-6 males contributed to each brood. This correlates with our behavioral observations that indicated multiple copulations between the female and several males in each experimental aquarium. A significant number of offspring from each brood were sired by non-sampled males (males that had copulated with females before capture) whose sperm had been stored by the female. This is the first direct evidence of multiple paternity in the Littorinidae. Results are discussed in reference to current theories of sperm competition, male precedence, and cryptic female choice.     

Inheritance of white spots on the shell of Littorina obtusata (Gastropoda, Prosobranchia)

The hypothesis on monogenic inheritance of white spot pattern on the shell of the gastropod snail Littorina obtusata was tested. Although in most cases our results did not contradict this assumption, the hypothesis cannot explain the observed segregation in general. An alternative hypothesis was suggested, according to which the presence of spot pattern is controlled by two complementary biallelic genes.     

Inheritance of longitudinal shell bands in the snails Littorina obtusata and Littorina saxatilis (Gastropoda, Prosobranchia)

The hypothesis of a monogenic inheritance of dark longitudinal bands on the shell in the gastropods Littorina obtusata and L. saxatilis was checked. One gene having two alleles proved to be responsible for the shell banding pattern in both of the species. The presence of bands was a dominant character in either case.     

Variation in populations of Littorina obtusata and L. mariae (Gastropoda) in the Severn Estuary

Populations of both species of flat periwinkle were examined at five discrete sites along the Severn Estuary, and at a marine site in Dyfed. The vertical distribution of the two species showed a clear partitioning of the shore – Littorina obtusata occupied the high/mid shore and L. mariae the low shore area. L. obtusata also extended further up the Severn Estuary than L. mariae. Maximum densities of both species were found in Milford Haven. Biometric investigations revealed intra and inter-specific differences in colour morphs, shell damage and size between populations. The possible influence of differential selection pressures at the various sites is discussed.     

Genetic differentiation in the molluscan species Littorina rudis and Littorina arcana (Prosobranchia: Littorinidae)

Six populations of Littorina rudis and ihree of L. arcana were screened for elcctrophorctically dctci table variation at 21 enzyme loci. These species reproduce by ovoviviparity and oviparily respectively, and both are members of the L. saxatilis species complex. The mean coefficient of genetic identity between the two species was high (0.957) and no diagnostic loci were observed. Allele frequency differences are maintained in sympatric populations of the two species. Intraspecific values of/ ranged from 0.944 to 0.995 and both species showed marked inter-population genetic heterogeneity, a feature consistent with their limited powers of dispersal. Heterozygosity is high in all populations of both species. Populations of I. rudis have higher mean heterozygosities (= 0.153) then L. arcane ( H = 0.132).     

Vertical migration and spawning inNodilittorina exigua (Gastropoda: Littorinidae)

The relation between the vertical migration in summer and spawning of a littoral-fringe periwinkle,Nodilittorina exigua, was studied by field observations and laboratory experiments. The snails moved down soon after the first appearance of their planktonic eggs in the sea and stayed at low shore-levels until early August. It was shown that the migration increased the chance of immersion of the snails, and therefore of releasing eggs by the females. Females immersed every day in the laboratory released more eggs than those which experienced some days of exposure to air, during the same period. The results obtained suggest the reproductive advantage of immersion at low levels. Nevertheless, the snails moved upward before the end of the spawning season. Typhoon swells increasing in frequency through the summer seemed to be responsible for the upward migration.     

On the taxonomy of Littorina africana (Mollusca: Gastropoda)

L. africana and L. knysnaensis are regarded as two morphs of a single species which exhibits a genetic cline along the south-eastern coast of southern Africa. The dark brown morph knysnaensis dominates the western, cooler end of the cline and is replaced by the pale blue morph africana at the warmer end of the cline. These conclusions are based on evidence from the latitudinal distributions, the complete range of intermediate forms regarding shell colour and shell morphology and the lack of differences in redular morphology, penial morphology or habitat.     

Field observations on the rhythmic up-and-down movement ofNodilittorina exigua (Gastropoda: Littorinidae)

Field observations on the activity pattern ofNodilittorina exigua were carried out in various tidal conditions in various seasons. The snails were stationary when dry on rock surface and began to move just after being splashed. On the rising tide they crawled upward aggregating in the awash zone but were not active around the time of high tide. On the receding tide they crawled predominantly downward, again in the awash zone. Position of the snails was high in spring tide and low in neap tide, changing in parallel to the change of high tide level. They were far more active in the awash condition than in the exposed or in the submerged conditions. For onset and termination of the movement, predominant influence of endogenous factors was suggested, except for onset of upward crawling. The distance of upward movement was larger when the high-tide level was higher, while that of downward movement was related to the time of high tide. Such movement patterns enable the snails to decrease the period of submergence and increase the period of their stay in the awash zone, and help them avoid long-term drying.     

Rhythmic egg-release in Littorina littorea (Mollusca: Gastropoda)

Batches of female Littorina littorea from four estuarine sites were kept in aquaria throughout their breeding season (December to July), some permanently submerged in circulating sea-water and others in a regime of simulated semi-diurnal tides of constant small amplitude. Females without the tidal experience released few eggs with little relation to the lunar month. Those in the tidal regime released more than four times the total number of eggs and released far more in periods of new and full moon than in periods of half-moon. Over the whole season, release was 1–5–3–25 times faster (at different sites) during full and new moon but the difference was more accentuated at the height of the season in February to early March and in early May. While individual females differed considerably and there were minor variations between the four sites, it is concluded that a two-weekly rhythm relates egg release to the spring-neap tide cycle and that this rhythm requires a tidal cycle of immersion and emersion for its maintenance. The special significance of such a rhythm for a species with pelagic eggs and larvae in an estuary is discussed.     

Species composition, comparative size and abundance of the genus Littoraria (Gastropoda: Littorinidae) from different mangrove strata along the East African coast

Variation in the abundance, distribution and size of four species of mangrove littorinid gastropods (genus Littoraria) was investigated using a nested sampling design at different spatial scales along the East African coast, from Tanzania to South Africa. Littorinid abundance and diversity decreased abruptly south of Inhaca Island at the southern end of the study area. All species presented a large-scale spatial variation in abundance, with L. subvittata showing the greatest abundance while L. intermedia was rare. Littoraria scabra and L. intermedia were found mainly at the seaward edge of the forests. Littoraria subvittata increased in abundance in the middle of the forest and towards the landward side. Littoraria pallescens occurred mainly at the seaward edge and in the middle areas in the Rhizophora zone. These small-scale variations show contrasting specific distribution patterns within the mangrove, likely reflecting different tolerances to physical factors and biological interactions. All species appeared decreased in shell height from north to south. Littoraria scabra was always significantly larger than other species at all mangrove study sites. Handling editor: P. Viaroli     

The breeding migration of Littorina arcana Hannaford Ellis, 1978 (Prosobranchia: Littorinidae)

Two species of rough winkle, the oviparous Littorina arcana and the ovoviviparous L. saxatilis are very similar in their outward appearance and are commonly syntopic. This has led some researchers to regard these species as conspecific and to suggest that reproductive modality is not a fixed characteristic. However, the syntopy of these species is much more complex than has been supposed and data presented here show how the seasonally reproducing L. arcana migrates downshore during its breeding period, into areas where L. saxatilis is comparatively sparse. This results in a separation of the breeding populations of the two species and consequently a spatial separation of their juveniles, at least in their early stages. Selection pressures on the breeding animals and young juveniles will therefore differ.     

Morphological variation in Lacuna parva (Gastropoda: Littorinidae) from different European populations

Shells of the littorinid gastropod Lacuna parva were compared from 23 European localities and postglacial deposits in Sweden. The shells from the recent and the postglacial populations are similar with the exception of the recent population from Ellekilde Hage, ?resund, Denmark. Shells from Ellekilde Hage are different in having especially well developed whorls and only one colour morph. Differences in life-cycle and radula morphometrics further distinguish the Ellekilde Hage population from populations from the Isle of Wight, UK, and Roscoff, France. No striking differences in penial morphology were observed between the populations. It is suggested that low salinity and subtidal occurrence might be the causative agents of the conchological differences exhibited by the ?resund population. Electronic Publication     

Microgeographic variation in allozyme and shell characters in Littorina saxatilis Olivi (Prosobranchia: Littorinidae)

Twenty-three enzymes and five shell parameters were screened in 11 subpopulations of Littorina saxatilis Olivi (= L. rudis Maton) occupying different habitats over a 1 km stretch of coastline. Shell morphology varied considerably and consistently with respect to degree of exposure, and since there is evidence that such morphology is at least partly under genetic control, it is likely that natural selection selects particular genotypes at particular locations. There was significant allozyme heterogeneity between neighbouring subpopulations, sometimes only metres apart, but little of the allozyme variability could be related directly to environmental pressures. Thus, with the exception of the Odh locus, the considerable morphological differentiation between snails from exposed and sheltered sites was not reflected in differentiation of those genes coding for electrophoretically assayed enzymes. At the Odh locus, virtually all the genetic differentiation between subpopulations was attributable to differentiation between habitat types. Two loci, Sod-1 and Aat-1, showed highly significant genetic disequilibrium, and possible reasons for this are explored. The population structure as assessed electrophoretically accords well with the stepping-stone model which permits greater differentiation of neighbouring populations than the island model, and which seems realistic in the ovoviviparously reproducing L. saxatilis, where the greater part of gene flow is likely to occur through the occasional migration of adults between contiguous populations.     

Intrasexual copulation and mate discrimination in a population of Nodilittorina radiata (Gastropoda: Littorinidae)

Intrasexual copulation and mate discrimination by Nodilittorina radiata (Gastropoda: Littorinidae) were studied on a concrete breakwater at Hakodate Bay, southern Hokkaido, Japan. Intrasexual (male–male) copulations were observed in 4.7–21.1% of copulating pairs on the shore. As females were relatively larger than males and males copulated with females larger than themselves, we hypothesized that males choose potential mates larger than themselves. However, two male mates showed no significant size preference in intrasexual copulations, suggesting that males do not choose relatively larger individuals as mates. In a laboratory mate-choice experiment, male N. radiata preferred to mate with females, indicating precopulatory sex identification. They copulated with males, however, at the frequency of 37%, perhaps because of sex misidentification.     

Shell colour polymorphism in a mangrove snail Littorina sp. (Prosobranchia: Littorinidae)

Shell colour polymorphism was examined in populations of a mangrove snail Littorina sp. in Queensland, Australia. Three morphs were recognized, yellow, red and brown, and morph frequencies varied both between widely spaced populations and between islands less than 1 km apart. Morph frequencies also varied with time of year. There was a relationship between shell colour and position on the tree, with yellow snails more often occurring amongst the foliage and brown snails more often on trunks and branches. In some populations yellow snails appeared to survive better than the other morphs, while in other populations there was no difference. The evidence for the maintenance of the polymorphism by natural selection is discussed.     

Locomotion of the filiform sperm of Littorina (Gastropoda,Prosobranchia)

Summary The filiform sperm of Littorina sitkana swims effectively in sea water and more viscous fluids, overcoming the problems of a non-uniform flagellar beat with an unusual mechanism, which involves three main events: (1) the sperm rotates anti-clockwise (looking from tail to head); then (2) stops rotating and stiffens itself to form a screw-shape, with the tail being held perpendicular to the middle piece, and finally; (3) reverses its rotation and propels itself forward in a clockwise spiral. The average velocity of sperm is approximately 185 ms with a rotational frequency of 24 revs. The mechanism of propulsion may involve two kinetic centers at opposite ends of the sperm, which coordinate their movements to produce anti-clockwise rotation, stationary twisting, or clockwise rotation, in a manner reminiscent of spirochaetes. Littorina sperm also exhibit slower methods of propulsion including swimming backwards (tail first) at 18 m, and gliding at about 30 m.The adaptive significance of the rapid propulsion is not obvious, because Littorina copulate and fertilize internally and at each stage in the transfer there are external aids to sperm transport, such as ciliary action (oviduct) and muscular expulsion (bursa and seminal receptacle). The filiform shape, however, is well-adapted for long-term storage in the female body. These points are discussed.     

Habitat and zonation patterns of Littoraria species (Gastropoda: Littorinidae) in Indo-Pacific mangrove forests

The zonation of 15 species of Littoraria in the scabra group is described, using a transect method, at 13 mangrove localities in Australia, Thailand, Malaysia and Hawaii. At these localities the littorinids occurred only on trees, and showed both vertical and horizontal patterns of zonation, with broad overlaps between co-occurring species. It is suggested that vertical distribution is determined mainly by behavioural responses, while the landward limits of horizontal distribution may be set by physiological tolerance. Several species occurred predominantly on either leaf or bark substrates, but no consistent associations with tree species were found. On the basis of local distribution patterns, Littoraria species can be classified as continental or oceanic in character. Oceanic species may be less tolerant of sediment in the water or on the substrate.     

Physical and biotic factors limiting the distribution and abundance of littoral molluscs on Macquarje Island (sub-Antarctic)

A number of factors have been examined for their influence on the limits of upper distribution and abundance of six species of adult molluscs in the rocky littoral zone of Macquarie Island. The molluscs were Amphineura: Plaxiphora aurata (Spalowsky), Hemiarthrum setulosum Dall; Gastropoda: Patinigera macquariensis (Finlay), Cantharidus (Plumbelenchus) coruscans (Hedley), Laevilittorina caliginosa (Gould), Kerguelenella lateralis (Smith), and the factors included habitat, temperature, desiccation, salinity, food, predation, and reproduction. Macquarie Island is mid-way between the southern temperate zone and the Antarctic and has an equable, oceanic climate. Because of differences in both geography and environmental stability, the results allow important comparisons to be made with those on the effects of factors limiting the distribution of littoral molluscs elsewhere. The effects of physical factors are similar to those examined in other regions as shown by tolerance to temperature, desiccation, and salinity. Habitat and zonal sequence of species are related to tolerances with the notable exception of the heat resistance of chitons which, when compared with that of gastropods, is greater than their distribution would suggest. In the field, the temperatures and desiccation of the animal bodies and the surrounding salinities do not reach critical lethal levels but the debilitating effects in combination with other adverse factors can be lethal — in particular, a combination of high temperatures and predation. Biotic factors have considerable influence on the distribution and abundance of the species and this may reflect the comparative stability of the physical environment. The aspects of the biotic factors which are important in this regard include morphology, mobility, food preferences, the presence of the large kelp Durvillea antarctica (Chamisso) Hariot, predation by birds and by isopods on enfeebled animals, and a brooding mode of reproduction. In all species, no single factor limits the upper distribution and factors vary in their importance for each species. Synergistic effects of factors are limiting and the effective combinations differ for each species. This stresses the danger of placing too much emphasis on results from studies of a single factor.     

Effects of environmental calcium deprivation on the egg masses of Physa marmorata Guilding (Gastropoda: Physidae) and Biomphalaria glabrata Say (Gastropoda: Planorbidae)

Eggs of the basommatophoran snails Physa marmorata and Biomphalaria glabrata were cultured in low concentrations of calcium to determine effects on growth and development. In both species there was some development in media with 0.12 mg/l Ca²⁺ but embryos were unable to hatch. 61.04% of embryos of P. marmorata could develop to hatching in 0.22 mg/l Ca²⁺ but those of B. glabrata required a level of 0.42 mg/l Ca²⁺, to attain even a 31.07% hatch. Marked effects on growth rate, embryo size and on time taken to achieve hatching were noted in both species at very low calcium levels. The possibility of cation-controlling mechanisms in the egg membrane is discussed.