Vicious circle in the intertidal: Facilitation between barnacle epibionts, a shell boring polychaete and trematode parasites in the periwinkle Littorina littorea

We studied interactions between three organisms associated with a common gastropod of northern Atlantic shores, the periwinkle Littorina littorea: barnacle epibionts Balanus crenatus, a shell boring polychaete Polydora ciliata, and tissue invading trematodes which use the periwinkles as first intermediate host. Snails collected shortly after barnacle settlement with > 50% cover of barnacles had significantly higher infestation of shell boring worms compared to unfouled snails, while trematode infestation was similar. The result was the same at two sites, and we conclude that the worm P. ciliata facilitates barnacle fouling on snails. The reverse was also the case. In an experiment with 14 weeks of exposure, snail treatments with barnacle epibionts had a significantly higher P. ciliata load than unfouled and cleaned snails. Again, trematode infestations were similar. The reciprocal positive interactions between barnacle epibionts and shell boring worms on snail houses is regarded as a case of facultative mutualism. On the other hand, for the snail basibiont, both barnacles and shell boring worms exert strongly negative effects by reducing fecundity, growth, and survival, resulting in a vicious circle for the snails. The combined effects of these associated organisms may rival in importance any competitive or predacious effects on the host L. littorea.     

An experimental investigation into the effect of crevice availability on abundance and size-structure in a population of Littorina rudis (Maton): Gastropoda: Prosobranchia

Littorina rudis (Maton) is a common species on a wide variety of rocky shores. At Newhaven L. rudis is generally abundant on boulders, juveniles being reliant upon empty barnacle shells for shelter. The experimental addition of crevices resulted in greatly increased population densities, suggesting that crevice availability is normally a limiting factor on this shore. A relationship is demonstrated between the size of artificially provided crevices and the mean shell lengths of the populations of L. rudis they support. The relevance of crevice availability and quality, including empty barnacle shells, to the abundance and size distribution of L. rudis and other littorines is discussed.     

Intraspecific variation in limblessness in vertebrates: a unique example of microevolution

Many species of arboreal marine snails in the genus Littoraria are polymorphic for shell colour, with morphs that correspond in both colour and frequency to those predominant in the habitat. Although the combined effects of selection for crypsis and apostasy have been suggested as the most likely reasons for this, they have not been demonstrated directly in the field. We investigated whether two parasitoid flies, Sarcophaga megafilosia and Sarcophaga meiofilosia, select for crypsis in Littoraria filosa . It was possible to compare the proportions that matched and did not match the background between samples of live and dead snails because the shells of individuals killed by these parasitoids, within which the larva and pupa develop, remain attached to the substratum. This comparison was necessary because these snails frequently move among different coloured microhabitats, which will tend to obscure any effect of selection for crypsis if only live individuals are censused. The method appeared reliable since there was no change in background colour and very little loss of shells between larviposition and emergence of flies. S. megafilosia killed a significantly greater than expected proportion of snails that did not match their background. In contrast, there was no evidence of selective attack by S. meiofilosia. These results are discussed in relation to previous work on polymorphisms in Littoraria species. We also suggest that the role of parasitoid insects as selective agents contributing to the maintenance of colour polymorphisms in terrestrial molluscs may have been overlooked.  © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 77, 367−377.     

Microgeographical variation in shell strength in the flat periwinkles Littorina obtusata and Littorina mariae

The strength of molluscan shells has been shown to vary in adaptive ways in a number of species and one of the main factors thought to be involved is shell-crushing by predators. A recent study found that the sibling species of flat periwinkle Littorina obtusata and Littorina mariae showed significant differences in the rates at which shell strength increased with shell length in specimens which had been collected from the same location, where the species were sympatric. This paper describes differences between the shells of the two species from a number of localities around Milford Haven in Dyfed, Wales, and local geographical variation in the shells. Littorina mariae, which is normally found at lower tidal levels than L. obtusata, matures at a smaller shell length. Both species reinforce the shell as they grow since shell strength, determined as the maximum force applied by a hydraulic tensile testing machine before the shell cracked, is strongly positively allometric; it increases at a rate close to the cube of shell length whilst isometric growth would result in strength increasing in proportion to the square of shell length. Because L. mariae matures earlier and reinforces the shell at a smaller size, the mature shell of L. mariae is substantially stronger on average than that of a similar sized but immature L. obtusata. At maturity the shell strengths of the two species are not very different despite the substantial difference in mean shell length. Strength varies significantly from shore to shore, and with the level of the shore from which the animals were collected. Strength increases down the shore in both species. Shell strength decreases with exposure to wave action in L. mariae but increases with exposure in L. obtusata; there is also substantial shore-to-shore variation which is not explained by exposure. Path analysis was used to explore the relationship between shell strength and other measured shell parameters (mass, length, height, thickness). The best predictor of shell strength in both species is a parameter which is heavily positively loaded on LN (shell mass) and strongly offset by negative loadings on LN (shell length) and LN (shell height). This is logical because for a given shell length a heavier shell will be thicker and stronger, whilst for a given shell mass a bigger shell will be thinner and therefore weaker. Such differential variation of shell mass and shell length explains most of the geographical variation observed in shell strength; shells are stronger in snails collected from one place than from another because, for the same shell length they are heavier or, to put it the other way, because at the same shell mass, they are smaller.     

The effects of exposure and predation on the shell of two British winkles

Changes in shell size and shell shape of the two British winkles Littorina nigrolineata and L. rudis were studied in relation to exposure and to crab-size. In both species, shells from exposed shores are smaller and more globose than those from sheltered shores. Also, in rudis of exposed shores the mouth is relatively wider. In shores of equally sheltered conditions, shells are bigger at those localities where crabs are large than at those localities where they are small. The largest shells are found in those localities where it is extremely sheltered, and the crabs are very large.
It is argued that on exposed shores, small shells are favoured because they have more possibilities than large ones to shelter in crevices and in barnacle interspaces, from the impact of winds and waves. A globose shell could accommodate more foot muscle and thus enable a stronger adherence to the rock; and an increased mouth diameter would increase the area of foot adherence to the rock. On sheltered shores, on the other hand, large, narrow-mouthed shells are favoured because they discourage crab predation, large crabs being abundant mainly on sheltered shores.
The possible significance of shell size and shape in relation to zonation is discussed, in view of the different predatory and physical conditions which prevail in different zones of the shore, and the different shell specializations which these conditions would require.     

Shell Morphological Variation of Littoraria angulifera among and within Mangroves in NE Brazil

The shell morphological variation of the periwinkle Littoraria angulifera (Lamarck, 1822) was studied in tropical northeast Brazilian mangroves. This area was selected because mangroves in different stages of regeneration, and thus different tree heights can be found. We evaluated whether differences in solar radiation due to differences in tree height influenced the distribution and shell morphology of L. angulifera, and carried out an experiment to test if individuals collected from mangroves with different tree heights differed in their resistance to desiccation. We also analysed if there were differences in L. angulifera shell length and shape between tidal levels within a mangrove. Finally, we tested if increased habitat complexity due to the presence of oysters could influence L. angulifera shell length and shape in different-statured mangroves. We predicted that the oysters could reduce desiccation stress on periwinkles in small-statured mangroves but have no effect in taller ones. Shell length and shape varied among mangroves, although a large variation within mangroves was also recorded. Shell proportionality (shell length:width ratio) increased with shell length, and this relationship differed among mangroves; however, no differences were found in a subsequent year. Individuals from small-statured mangroves survived longer than those from taller mangroves in the desiccation experiment, with a weak correlation between shell proportionality and loss of mass. The presence of oysters had no apparent influence on shell morphology that could be correlated with reduced desiccation stress. The patterns found varied greatly both at small and large spatial scales, suggesting that future studies should evaluate phenotypic and genetic variation at the same time to properly understand variation in L. angulifera shell morphology.     

Contrasting reproductive traits in two species of mangrove‐dwelling littorinid snails in a seasonal tropical habitat

Previous studies on the reproductive biology of littorinid snails have focused on rocky shore species, investigating how these gastropods can achieve maximal reproductive success, as well as on processes of sexual selection. This study documented differences in the reproductive traits of two mangrove‐dwelling littorinids, Littoraria ardouiniana and L. melanostoma, in Hong Kong. Reproductive activity of both species was most intense during the summer months. Mating pairs of the two species generally occurred in the tree canopies. Few false mating pairs (same sex or heterospecific pairs: <10%) were recorded, and members of both species showed size‐assortative mating. Littoraria ardouiniana had a shorter reproductive season but a higher intensity of mating and higher seasonal fecundity, than did L. melanostoma. Members of both species showed bi‐lunar periodicities of egg or larval release, synchronized with spring tides. Fecundity showed a strong positive relationship with body size in L. ardouiniana, but not in L. melanostoma. Females of L. ardouiniana released entire broods of larvae in a single brief event, whereas females of L. melanostoma released fewer eggs over 1–8 d. Release of larvae in L. ardouiniana involved a series of short bursts and was much faster than the trickle release of eggs in L. melanostoma. The contrasting reproductive traits in these two species represent different strategies to optimize reproductive success in mangrove habitats.     

Divergence in the shell morphology of the land snail genus Aegista (Pulmonata: Bradybaenidae) under phylogenetic constraints

The role of natural selection in phenotypic evolution is central to evolutionary biology. Phenotypic evolution is affected by various factors other than adaptation, and recent focus has been placed on the effects of phylogenetic constraints and niche conservatism on phenotypic evolution. Here, we investigate the relationship between the shell morphology and habitat use of bradybaenid land snails of the genus Aegista and clarify the causes of the divergence in shell morphology among phylogenetically related species. The results of ancestral state reconstruction showed that arboreal species have evolved independently from ground‐dwelling species at least four times. A significant association was found between shell shape and habitat use, despite the existence of a certain degree of phylogenetic constraint between these traits. A principal component analysis showed that arboreal species tend to have a relatively high‐spired shell with a narrow umbilicus. By contrast, ground‐dwelling species have a low‐spired shell with a wide umbilicus. Although the latitude and elevation of the sampling locations showed no relationship with shell morphology, the geology of the sampling locations affected the shell size of arboreal species. The development of a well‐balanced shell shape is one effective method for reducing the cost of locomotion under the force of gravity in each life habitat, resulting in the divergence in shell morphology and the independent evolution of morphologically similar species among different lineages. The present study suggests that ecological divergence is probably the cause of shell morphology divergence in land snails. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 114 , 229–241.     

Mechanisms and consequences of shell fouling in the kelp snail,Norrisia norrisi (Sowerby) (Trochidae): Indirect effects of octopus drilling

The kelp snail, Norrisia norrisi (Sowerby), dwells on the large kelps Macrocystis pyrifera (L.) and Eisenia arborea Areschoug, and is rare on benthic substrata. Approximately 4% of the snail population is dislodged from plants each day. Per capita mortality of snails on the benthos is an order of magnitude greater than individuals on kelp. Even though snails displaced to the bottom actively seek a plant to climb, we found that between 20–40% of marked snails released on the bottom were killed over a 24-day period. While most N. norrisi are free of epibionts on their shells, ≈30% of the population are encrusted by the barnacle Megabalanus californicus (Pilsbry). The majority of barnacle fouling was the result of failure by Octopusbimaculatus (Verrill) to kill Norrisia norrisi; barnacle cyprids settled in octopus drill holes in the shells of live kelp snails. N. norrisi encrusted by barnacles were less effective in exploiting kelp compared with unfouled snails. Moderately to heavily fouled snails were dislodged from kelp twice as frequently and remained on the benthos much longer than snails without epibionts. Laboratory tests revealed that escape velocities of N. norrisi from asteroid predators were much reduced by shell fouling, and that snails carrying barnacles were most likely to be captured and consumed. Hence octopus increase mortality of kelp snails in two ways. First, octopus can kill N. norrisi encountered on the bottom. Second, kelp snails drilled by octopus, but not killed outright, will become fouled by barnacles. This results in a subsequent risk of mortality from all benthic predators that is at least 8-fold greater than for unfouled kelp snails.     

The effects of predation by the mummichog, Fundulus heteroclitus (L.), on the abundance and distribution of the salt marsh snail, Melampus bidentatus (Say)

Enclosure and exclosure experiments were conducted in Canary Creek marsh to examine how predation by a killifish, Fundulus heteroclitus (L.), affects the abundance and size distribution of the salt marsh snail, Melampus bidentatus (Say). Enclosures (7.6 × 19.7 m) were stocked with Fundulus heteroclitus at densities of one-half normal, normal, twice normal, and four times normal density. Fish exclusion pens were also built. In both years of study, the mean density of snails increased significantly in pens where fish were excluded or their density reduced. During the same period in each year, the density of snails in pens containing higher than normal fish density fell by ≈50%.

Fish density also affected the size distribution of snails within pens. In both years, mean shell length of snails in the pen with the highest density of fish was significantly greater, and mean shell length of snails within fish exclusion pens was significantly lower than in all other treatments. Gape size limitation of F. heteroclitus causes selective predation on small snails and apparently is responsible for the difference in mean shell length among treatments.

Density and size distribution measurements of the natural Melampus bidentatus population in Canary Creek marsh were also taken. Larger snails were found to occur in the low marsh zone, whereas smaller snails occurred in the high marsh zone. Since the high marsh area is flooded less often than the low marsh, and the grass types found in the high marsh zone are known to afford protection from fish predation, the distribution of snails in the marsh is consistent with the idea that fish predation is an important factor influencing the distribution of snails in Canary Creek marsh.     

Crayfish induce a defensive shell shape in a freshwater snail

Abstract. This study was undertaken to determine whether intraspecific variation in shell morphology of the freshwater snail Elimia (= Goniobasis ) livescens is caused by predator-induced morphological changes. Juvenile snails from 3 populations were grown in the presence of effluent from predatory crayfish feeding on conspecific snails or in effluent from conspecific snails only. Snails from one population, Clear Creek, exhibited a predator-induced morphology; they grew a narrow body whorl when exposed to the effluent from crayfish. Experimental feeding trials with crayfish and snails from Clear Creek were conducted to determine whether a narrow body whorl reduced predation in the presence of the crayfish Orconectes propinquus . In the feeding trials, snails with a narrow body whorl were eaten less frequently than snails with wider shells. However, there was no difference in overall size (length) between snails that were eaten and those that were not. Thus, juvenile snails from Clear Creek exposed to the presence of crayfish were induced to develop relatively narrow shells, which reduce the risk of successful attack by these crayfish. Hence the induced shell morphology is probably a defense against predation by crayfish.     

Resource allocation,demography and the radiation of life histories in rough periwinkles (Gastopoda)

Applicability of life-history theory to higher levels of comparison (from populations, through ecotypes to sibling species) was investigated in rough periwinkles, whose life histories have diversified since colonization of the North Atlantic by an oviparous ancestor in the upper Pliocene. Comparisons were made among populations of the ovoviviparous Littorina saxatilis, between L. saxatilis and its ecotype, L. neglecta (with an annual life history) and between the sibling species L. saxatilis and L. arcana, the latter of which retains the ancestral oviparity. Resource-allocation priority, reproductive effort and related trade offs were compared between the ecotypes and the sibling species by measuring changes in flesh mass and reproductive output in snails subjected to different degrees of food deprivation, and by measuring mortality rate of snails stressed by desiccation, high temperature and low salinity. Body size had a marked effect on all parameters, but after statistically removing this effect there remained no significant differences in allocation among ecotypes or species. Published demographical data were reviewed for correlations between habitat, mortality regime and life-history characteristics. Populations of L. saxatilis varied principally in size at birth and in adult size. Theoretical premises based on density-dependent versus density-independent mortality regimes could not explain these trends. Instead, size at birth may have reflected the mechanical, physiological or biological nature of mortality risk rather than its density dependence or independence. Adult size reflected the available sizes of crevices used for shelter and perhaps also the quality of feeding conditions. Radiation of life histories within the rough periwinkles is interpreted as a series of adaptations to a progressively wider range of habitats. The transition from oviparity to ovoviviparity allows colonization of estuaries, saltmarshes and pebble beaches too hazardous for naked egg masses. The transition from a perennial to an annual life history in barnacle ecotypes follows from allometric re-scaling of morphological and physiological parameters, enabling reproduction and brooding to occur at the small body size necessary for life within empty barnacle tests. This suite of adaptations allows exploitation of a relatively benign microhabitat that occurs almost ubiquitously on exposed rocky shores of the temperate North Atlantic. The persistence of oviparous forms, presumably in the face of competition from sympatric ovoviviparous forms, remains unexplained.     

Modification of responses of newly hatched snails by exposure to odors during development

Predatory marine snail larvae and embryos were exposed to preyodors (oyster, mussel and barnacle) during development. Whenjuvenile snails hatched they were tested by bioassay to determineeffects of this prior odor exposure. Juvenile snails were testedwith oyster, mussel and barnacle odors and with partially purifiedbarnacle odor of known attractiveness. Independent of priorodor exposure, only solutions containing barnacle odor wereattractive. Snails exposed during development to prey odorsnot in themselves attractive were more responsive to barnacleodors than snails that developed in barnacle odors. Snails notexposed to prey odors during development were intermediate intheir responsiveness. Field bioassays showed detectable attractantlevels in an environment containing barnacles. Attractant activitywas not detected in an environment where barnacles were absent.Responses of snails that developed in field odor conditionswere similar to those of snails that developed in the presenceof barnacles. Odor exposure during development is importantin determining future responses of these predatory snails toprey odors. ¹Duke University Marine Laboratory, Pivers Island, Beaufort,NC 28516, USA ²University of Miami RSMAS/MAC 4600 Rickenbacker Causeway, Miami,FL 33149, USA ³Suffolk University, Department of Biology, Beacon Hill, Boston,MA 02114, USA     

Shell shape and land-snail habitat in a Mediterranean and desert fauna

The bimodal distribution of shell shape (height: diameter), that is found in various geographically widely separate and taxonomically distinct land snail faunas of many different regions of the world, occurs also in a Mediterranean fauna and in a desert fauna that is derived from it. The desert fauna is, however, closer to the bisector than the Mediterranean one. High-spired snails are mainly rock-dwellers, and equidimensional to low-spired snails are bush-dwellers or soil-diggers, with a few rock-dwellers; litter-dwellers are small-sized species that may have either high- or low-spired shells. These results are discussed in adaptive terms. Litter is probably the more primitive of these micro-environments. Many of the small, litter-dwelling snails are ovo-viviparous rather than oviparous, perhaps so as to avoid attacks on the eggs by saprophytic fungi. The shift away from the litter environment is accompanied by a trend to abandon the ovo-viviparous strategy, in favour of oviparity, the snail using its foot to dig into the soil and lay eggs. The conchometric differences between bush-, ground- and rock-dwelling snails may perhaps reflect selective pressure to increase the size of the foot; and constraints of a habitat that consists of narrow interspaces between rocky boulders. Snails that habitually dig into the ground during periods of inactivity, and roam over the ground when active, requires a very large foot and, consequently, a very large-mouthed shell to accommodate it; the result is an equidimensional shell, globose or turbiniform in shape. Snails that climb up vertical vegetation would also require a large foot, and consequently a large-mouthed shell to contain it. A fully globose shell would however be disadvantageous, since it might cause undesired torque. Hence, bush-dwellers tend to be flatter than soil-diggers. Snails that habitually live in rock crevices, and on hard substrata, would not require a very large foot; they would need a narrow shell, both to enable easy manoeuvring through crevices and to reduce torque, the result being a small-mouthed, usually high-spired shell. The classification of land snails into bush-, soil- or rock-dwellers closely follows the taxonomic classification. In those species that depart from the habitat that is typical of their taxonomic group towards another habitat, the shell alters its shape accordingly.     

Phenotypic homogeneity of two intertidal snails across a wave exposure gradient in South Australia

Dislodgement by the large drag forces imparted by breaking waves is an important cause of mortality for intertidal snails. The risk of drag-induced dislodgement can be reduced with: (1) a smaller shell of lower maximum projected surface area (MPSA); (2) a streamlined shell shape characterized by a squatter shell; and/or (3) greater adhesive strength attained through a larger foot area or increased foot tenacity. Snails on exposed coasts tend to express traits that increase dislodgement resistance. Such habitat-specific differences could result from direct selection against poorly adapted phenotypes on exposed shores but may reflect gastropod adaptation to high wave action achieved through phenotypic plasticity or genetic polymorphism. With this in mind, we examined the size, shape and adhesive strength of populations of two gastropod species, Austrocochlea constricta (Lamarck) and Nerita atramentosa (Reeve), from two adjacent shores representing extremes in wave exposure. Over a 5 day period, maximum wave forces were more than 10 times greater on the exposed than sheltered shore. Size-frequency distributions indicate that a predator consuming snails within the 1.3-1.8 cm length range regulates sheltered shore populations of both snail species. Although morphological scaling considerations suggest that drag forces should not place physical limits on the size of these gastropods, exposed shore populations of both snails were small relative to the maximum size documented for these species. Therefore, selective forces at the exposed site might favour smaller individuals with increased access to microhabitat refuges. Unexpectedly, however, neither snail species exhibited between-shore differences in shape, foot area or foot tenacity, which are likely to have adaptive explanations. Hence, it is possible that these snails are incapable of adaptive developmental responses to high wave action. Instead, the homogeneous and wave-exposed nature of Australia's southern coastline may have favoured the evolution of generalist strategies in these species.     

Small- and broad-scale patterns of distribution of the upper-shore littorinid Nodilittorina pyramidalis in New South Wales

Abstract mall-scale (within-shore) and broad-scale (among-shore) patterns of distribution, abundance and size structure of Nodilittorina pyramidalis were measured on a number of shores in New South Wales. This species showed significant patchiness in distribution and size structure. The mean density and mean shell length of snails differed among patches of shore only a few metres apart, among different heights on the shore and from shore to shore. Differences in densities appeared to be strongly correlated to the distribution of particular microhabitats, especially cracks, crevices and pits within the rock surface. This relationship and the presence or absence of these microhabitats appeared to determine the patterns of distribution of this species within and among different shores. The sizes of snails were not correlated with densities of Nodilittorina nor with the coexisting species, Littorina unifasciata, in the populations examined. Processes influencing the distribution and abundance of Nodilittorina were not investigated, but the patterns of distribution found on these shores suggest that recruitment was limited to areas of topographic complexity, ft also seems likely that limited dispersal by the adult snails retain them within restricted patches on the shore.     

Predator-induced defenses in a salt-marsh gastropod

Predator-induced defenses are among the most ecologically important forms of phenotypic plasticity. Although predation and induced defenses are well documented in rocky-intertidal systems, they have received less attention in soft-bottom communities. Shell-crushing predators are common in soft-bottom, vegetated habitats, which often exhibit substantial spatial heterogeneity in predation intensity. We examined variations in shell morphology of the salt-marsh periwinkle, Littoraria irrorata, among marsh microhabitats in the northern Gulf of Mexico that vary in their accessibility to predatory blue crabs, Callinectes sapidus. Littoraria from high-predation sites exhibited more extensively calcified apertural lips and narrower apertural openings relative to snails from low-predation sites. Thick apertural lips generally increased the handling time required by Callinectes to breach Littoraria shells in laboratory experiments, although the method of shell entry used by crabs was dependent on the crab:snail size ratio. Apertural-lip thickness was not related to past predation events in field-collected snails. Snails exposed to water treated with the effluent of Callinectes and crushed conspecifics produced significantly thicker apertural lips than controls, with a response time and morphological extent comparable to that of their rocky-shore counterparts. This study underscores the widespread occurrence of predator-induced plasticity in marine gastropods and emphasizes its role in soft-bottom, vegetated marine habitats, where shell-crushing predation can be as prevalent a selective force as in the rocky intertidal.     

Age-dependent zonation of the periwinkle Littorina littorea (L.) in the Wadden Sea

On sedimentary tidal flats near the island of Sylt (German Bight, North Sea) abundance and size distribution of periwinkles, Littorina littorea L., were studied in low intertidal and in shallow and deep subtidal mussel beds (Mytilus edulis L.). In low intertidal mussel beds, surveys revealed that high densities (1,369±571 m^–2) of juvenile snails (≤13 mm) were positively correlated with strong barnacle epigrowth (Semibalanus balanoides L. and Balanus crenatus Bruguière) on mussels. A subsequent field experiment showed that recruitment of L. littorea was restricted to the intertidal zone. Abundances of periwinkles (213±114 m^–2) and barnacles abruptly decreased in the adjacent shallow subtidal zone, which served as a habitat for older snails (>13 mm). L. littorea was completely absent from disjunct deep (5 m) subtidal mussel beds. Snail abundance varied seasonally with maxima of >4,000 m^–2 in low intertidal mussel beds in October and minima in July, just before the onset of new recruitment. I suggest that the presence of cracks and crevices among the dense barnacle overgrowth in intertidal mussel beds favoured recruitment and survival of juvenile snails. Larger (older) specimens are assumed to actively migrate to the less favourable adjacent subtidal. Therefore, intertidal mussel beds are considered as nurseries for the population of L. littorea in the Wadden Sea. Received in revised form: 25 September 2000 Electronic Publication