Shell colour polymorphism associated with substrate colour in the intertidal snail Littorina saxatilis Olivi (Prosobranchia: Littorinidae)

Littorina saxatilis Olivi (1792), the rough winkle, is highly polymorphic in shell colour. Shell colour frequencies were studied at six locations in south-western Wales, U.K., each at a geological contact between red sandstone and grey limestone or volcanic rock. At each site shell colour frequencies were determined in samples from the contact zone and on red or grey rock on either side. Highly significant associations were found between shell colour frequencies and substrate colour. Grey shells were always more common on grey rock than on red rock, and brown shells were usually more common on red than on grey rock, suggesting selection for cryptic colouration. Shell colour frequency differences were also found between replicate samples taken only 5 m apart from the same kind of rock, and between samples from the same kind of rock at the six study sites. These latter differences suggest that selection for camouflage is not the only factor involved in maintaining shell colour polymorphism in this species.     

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.     

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.     

Shell colour variation in Bullia digitalis, a sand-dwelling, intertidal whelk (Gastropoda: Prosobranchia)

Bullia digitalis is an intertidal whelk that lives on sandy beaches in South Africa. It is highly variable in shell colour, with individuals varying from white to dark brown. This paper describes shell colour variation of B. digitalis at seven sites, along a 230 km coastline east of the Cape Peninsula. Seven colour forms were found: striped, violet, banded violet, banded brown, orange, pale yellow and white. These forms are probably genetically determined morphs. The striped form is the most common at all sites, constituting 53–62% of each sample. The violet is the second most common morph. Its frequencies are remarkably stable at 15–17%. The striped form blends well into the sandy environment and may therefore be of considerable cryptic value in concealing B. digitalis from predators. The violet form is highly conspicuous. Its stable frequency throughout the study area may represent a genetic balance that is not relevant to any visual advantages of the violet colour.     

The role of selection in the colour polymorphism of Littorina rudis Maton and Littorina arcana Hannaford-Ellis (Prosobranchia: Littorinidae)

Forty-one mixed samples of winkles containing the closely related species, Littorina rudis and L. arcana , were collected from different parts of the British Isles. Littorina rudis was the more ubiquitous species, with L. arcana being more-or-less confined to vertical cliffs and rocks. The frequencies of different shell colour patterns were determined for both species in each sample. Several colour morphs were diagnostic of one or other of the species over large areas, which confirms that these are separate species. The frequencies, in the two species, of the two commonest morphs, brown and fawn, were strongly correlated; in several other morphs their presence and absence in the two species was significantly associated; the levels of phenotypic diversity in the two species were also correlated. These patterns could not have arisen if the variation between shores was the result of genetic drift or founder effects, so some form of selection is implicated. In the brown morph there is some evidence for frequency-dependent selection, and it is suggested that the polymorphism may be maintained by visual predators through apostatic selection.     

Mate discrimination in Littorina littorea (L.) and L. saxatilis (Olivi) (Mollusca:Prosobranchia)

The ability of males of Littorina littorea and L. saxatilis to discriminate between mates of different sex, species and size was examined. In partner choice experiments males of L. littorea had the possibility to initiate a copulation with either a female or a male. The males did not show a preference for either sex. There was therefore no evidence that they could determine the sex of a conspecific prior to copulation. The duration of intrasexual copulation was considerably shorter than for intersexual copulation, both in the field and in laboratory experiments. For the two species, intersexual copulations were far more frequent than intrasexual ones. This can partly be explained by the difference in copulation time.Few interspecific copulating pairs were found on the shore. This may reflect a low interspecific encounter rate rather than a mechanism of species recognition. On all of these occasions, however, the active male was of L. saxatilis. It is argued that selection against precopulatory species and sex recognition is a more likely explanation than an hypothesis that states that the required mutations for precopulatory mate identification has not yet occurred. L. littorea males copulated longer with large than with small females. Copulation time was short with parasitized females, which are sterile or of low fecundity. The allocation of mating effort by males is discussed.     

Habitat related allozyme variation on a microgeographic scale in the marine snail Littorina mariae (Prosobranchia: Littorinacea)

The marine snail Littorina mariae Sacchi & Rastelli occurs in high numbers in the littoral zone on fucoid macro-algae. The eggs are laid on the seaweed and d e velopment to miniature snails takes place without any pelagic larval stage. We have mapped the genetic variation of 30 enzyme loci in populations from eight small islands within 15 km of each other on th swedish west coast. The original intention was to investigate the magnitude of gene flow within and between islands. However, we soon realized that our basic assumption of neutral genetic variation was forcefully violated in at least one locus, arginine kinase ( Ask ). Allele frequencies of Ark were strongly associated with type of habitat. Therefore the main part of this study focused on allele frequency distribution in different habitats. Eight of the 30 loci screened were polymmorphic but we mainly considered the four most polymorphic ones (total heterozygosity between 0.32 and 0.57). All four showed significant heterogeneity between subpopulations, Ask especially so; 42% of the total variation of Ark was explained by differentiation between samples, and 90% of this variation was attributed to differences between different types of habitats (more or less exposed to wave action). In contrast, peptidase ( Pep-1 ) and phosphoglucomutase ( Pgm-2 ) varied in a way predicted by neutral theory; between sample variation being mainly attributed to differentiation bvetween islands. The variation in phosphoglucose isomerase ( Pgi ) was less consistent. In some islands there w as an obvious difference between different habitats, but on other islands we found no significant difference.Not taking into account the extreme, presumably sclected, variation in Ark , we concluded that the metapopulation of Littoriana mariae that we studied was divided into semi-isolated populations, between which the average rate of migration was in the range of a fe w individuals per generation.     

Sexual selection and non-random mating for shell colour in a natural population of the marine snailLittorina mariae (Gastropoda: Prosobranchia)

In order to estimate the three independent components of mating behaviour, sexual selection in females, sexual selection in males and mating pattern, we studied the distribution of shell colour morphs among mating pairs and between copulating and non-copulating snails in four subsamples of a natural population ofL. mariae. The colour of the shell, the sex and a qualitative estimate of age was recorded for every snail. We found sexual selection acting against one of the two commonest colours (yellow) among the young females. However, in males none of the eight shell colour morphs was favoured during matings. Male sexual choice or differences in female sexual activity may cause the sexual fitness disadvantage of yellow females. Moreover, individuals of different colour morphs did not mate at random, rather dissasortatively. A behavioural choice among shell colour morphs or a non-random microdistribution of the morphs may cause the departure from random mating in this population.     

Observations with a different approach to morphological colour variation in an isopod population (Crustacea: Isopoda)

Morphological colour variation in Idotea baltica basteri Audouin, 1827 (Isopoda: Crustacea) is reported for the first time on populations from a brackish water lake (Kucukcekmece Lagoon) in Turkey. According to pigmentation characteristics, the isopods were described and sorted to ten different colour morphs which are named as maculata, maculata-lineata, uniformis, uniformis-lineata, immaculatum, immaculatum-lineata, nigrum, nigrum-lineata, albafusca, and albafusca-fasciatum. The similarity of the morphs was compared by defining digit codes to some of the visual pigmentation characteristics and calculating the Bray-Curtis similarity index of the colour morphs. The albafusca and albafusca-fasciatum forms were found to be the most similar to each other (similarity 96%), whereas immaculatum forms were considered the most different from maculata-lineata (similarity 50%). The morphs named maculata and maculata-lineata, having the highest abundance with 70%, belong to the most successful forms which may achieve crypsis by background matching in their living habitat in Kucukcekmece Lagoon which has sandy and rocky bottom surface area.