Land Snail Cepaea Research Articles

Fine mapping of the Cepaea nemoralis shell colour and mid-banded loci using a high-density linkage map

Molluscs are a highly speciose phylum that exhibits an astonishing array of colours and patterns, yet relatively little progress has been made in identifying the underlying genes that determine phenotypic variation. One prominent example is the land snail Cepaea nemoralis for which classical genetic studies have shown that around nine loci, several physically linked and inherited together as a ‘supergene’, control the shell colour and banding polymorphism. As a first step towards identifying the genes involved, we used whole-genome resequencing of individuals from a laboratory cross to construct a high-density linkage map, and then trait mapping to identify 95% confidence intervals for the chromosomal region that contains the supergene, specifically the colour locus (C), and the unlinked mid-banded locus (U). The linkage map is made up of 215,593 markers, ordered into 22 linkage groups, with one large group making up ~27% of the genome. The C locus was mapped to a ~1.3 cM region on linkage group 11, and the U locus was mapped to a ~0.7 cM region on linkage group 15. The linkage map will serve as an important resource for further evolutionary and population genomic studies of C. nemoralis and related species, as well as the identification of candidate genes within the supergene and for the mid-banding phenotype.

Shell colouration and different introductions of the land snail Cepaea hortensis (Gastropoda: Helicidae) into Western Ukraine

The primary introduction of C. hortensis into Western Ukraine, most likely in the second half of the 20th century, led to the formation of many populations characterised not only by a reduced phenotypic composition, but also by a peculiar phenotypic marker (absence of banded shells with a non-white ground colour). White banded shells, common in Western Ukraine, are also found in different parts of the present species range, but less frequently than yellow banded ones. When studying the shell colour and banding polymorphism in C. hortensis, white shells are often combined into one group with yellow ones. Therefore, the findings of these two phenotypes as well as white unbanded shells in Europe were mapped based on the analysis of more than 7 thousand photographs from citizen science database iNaturalist. The results of later introductions of C. hortensis to Western Ukraine, which may occur through different garden centres, are easily distinguished by the presence of yellow banded and/or pink shells. And the presence of a dark lip in all pink shells can be a phenotypic marker of snails, whose ancestors were introduced to the west of Ukraine through the garden centre “Club of Plants” near Lviv. Only near this garden centre were also found three brown-shelled snails not known for other introduced Eastern European populations of C. hortensis.

Morph‐dependent nematode infection and its association with host movement in the land snail Cepaea nemoralis (Mollusca, Gastropoda)

AbstractHost behaviour can be influenced by parasitic risk and infection through a variety of direct and indirect mechanisms. We can expect individuals expressing different phenotypes to also differ in the ways their behaviour is altered by parasites. We used the land snail Cepaea nemoralis, a species with various shell colour morphs differing in behaviour and physiology, as a model to study the link between parasite response and individual behaviour variation. We analysed metazoan parasite abundance and its relation to behaviour (movement and food intake) in snails from three shell morphs (from light unbanded to darker five‐banded) and from two neighbouring populations from contrasting environments. Snails were parasitized by mites, trematodes, and nematodes, from rarest to most frequent. We confirm that terrestrial gastropods can defend against infection by trapping parasitic nematodes in their shell. We show that nematodes encapsulated in shells can uncover past infections even when a snail population is currently nematode‐free, possibly due to parasite seasonality. We present new observations suggesting that not only nematodes but also mites might be encapsulated in shells. Infection levels varied between morphs, with darker snails harbouring fewer nematodes. Behaviour (movement and food intake) was linked to the nematode, but not trematode infection. Individuals with higher nematode load ate less, irrespective of morph and population. The most‐infected morph (unbanded snails) showed reduced activity in the nematode‐infected population compared to the one that was nematode‐free at sampling time. As a result, parasites may cancel previously described behavioural differences between morphs. We discuss the possible mechanisms explaining morph‐dependent responses to parasites, and how parasite risk may be an unseen force shaping C. nemoralis morph variation in space and time. We conclude with the possible ecological consequences of the link, mediated by shell colour, and between thermal and immune responses.

Comparative analysis of phenotypic variability of introduced land snail Cepaea nemoralis (Gastropoda: Helicidae) in two large Eastern European cities

Variation in the shell colour and banding polymorphism of Cepaea nemoralis was recorded in 20 sites in Minsk, Belarus (a total of 3965 adults collected in 2014–2021). This variation was compared with that in 16 sites from Lviv, Western Ukraine (total 3235 adults collected in 2019–2021). Unlike in Lviv, a remarkable spatial differentiation of the phenotypic composition was found in Minsk. The samples collected in the north-eastern part of Minsk were characterized by a greater degree of phenotypic diversity and by higher frequencies of unbanded and brown shells. Samples from the southern and eastern parts of the city were generally lighter and characterised by high frequencies of shells with a single central band (mid-banded). Differences between the two parts of Minsk were greater than those between Minsk and Lviv. This may be related to the history of colonization of Minsk by C. nemoralis. The Lviv samples were, on average, darker, they contained less frequently mid-banded shells and more frequently unbanded shells, especially pink unbanded ones. The Fst values characterizing the level of phenotypic variability and calculated for Minsk and Lviv are quite high and comparable with those for other urban areas recently colonized by C. nemoralis.

Level of oxidative stress for the land snail Cepaea nemoralis from aged and bioremediated soil contaminated with petroleum products.

Here, we investigated whether the widely distributed snail Cepaea nemoralis could be used as a suitable sentinel animal for assessing the effects of soil contaminants-petroleum oil derivatives-after years of soil ageing and treatment with a bacterial formulation. Oxidative stress was assessed in the foot and hepatopancreas of C. nemoralis L. exposed to soil contaminated with unleaded petrol, spent engine oil or diesel oil and bioremediated with a bacterial formulation (soil was used 2years after contamination and bioremediation process). We measured total antioxidant capacity, catalase and glutathione transferase activity and concentrations of superoxide anions, hydrogen peroxide and protein carbonyls in the foot and hepatopancreas of snails after 2 and 4weeks of treatment. The studied antioxidant responses appeared largely to be tissue and remediation process specific, while the concentrations of superoxide anions, hydrogen peroxide and protein carbonyls depended on time of exposure, tissue type and the type of contaminants, but mostly not on the remediation process. Generally, changes in the concentrations of superoxide anions, hydrogen peroxide and protein carbonyls in the hepatopancreas of snails seemed to be a suitable measure to assess the risk of animals exposed to soil contaminated with petroleum substances and used after many years of ageing and treatment with a microbial formulation.

Variability of the Phenotypic Composition of Cepaea hortensis (Gastropoda, Helicidae) in Western Ukraine: in Space and Time

The variability of the phenotypic composition of the introduced land snail Cepaea hortensis was analyzed in 6 groups of samples (in total, more than 31 thousand specimens), differing in spatial location or collection time. The results obtained confirmed the considerable uniformity of this composition, which has not yet been significantly influenced by the relatively recent repeated introductions of C. hortensis associated with the activity of garden centers and contributing to an increase in phenotypic richness at separate, still very spatially limited sites. The Western Ukrainian populations of C. hortensis are characterized by low values of the inbreeding coefficient, caused by a common origin and quite possibly by climatic selection favoring unbanded shells.

Introduced land snail Cepaea nemoralis (Gastropoda: Helicidae) in Eastern Europe: spreading history and the shell colouration variability

The available data on the present distribution of Cepaea nemoralis in Ukraine, Belarus and the European part of Russia, the history and possible ways of penetration of this species into various parts of Eastern Europe, as well as the colour and banding polymorphism of its shells are analyzed. It has been suggested that the process of rapid spreading of C. nemoralis in the three compared Eastern European countries began at approximately the same time (late 20th – early 21st centuries) and that it may be caused by active and practically uncontrolled importation of seedlings from other European countries, as well as by global climate change, which can contribute to better survival of snails outside their natural range. Despite the possible initial limitation of the phenotypic and genetic diversity associated with the accidental transfer of a limited number of individuals, in the Eastern European colonies, in general, a fairly large variety of the inherited traits of the shell colouration remains. This concerns, first of all, the polymorphism in the shell ground colour (yellow, pink, less often brown) and the main variants of the shell banding (unbanded, mid-banded, five-banded, less often three-banded).

The first records of an invasive land snail <i>Cepaea nemoralis</i>(Stylommatophora: Helicidae) in Central and Southern Ukraine

Two colonies of an invasive West European land snail Cepaea nemoralis are reported from Kyiv region of Ukraine, as well as a third locality where snails were present in 2019, but no living individuals were found in 2020. One more locality is reported from Odessa City in Southern Ukraine. These localities are the most south-eastern ones for the range of C. nemoralis and are the first known records of this species in Ukraine outside its western regions, with closest previously published confirmed colonies in more than 400 km. Distribution of this species in Eastern Europe is discussed, it is argued that invasion ofC. nemoralis is probably driven by the same processes as invasion of the Spanish slug, Arion vulgaris, that invaded Eastern Europe during the last decades.

Intrapopulation variation in shell morphological traits and banding polymorphism of the land snail Cepaea vindobonensis (Gastropoda; Pulmonata; Helicidae)

The main goal of this paper was to analyze intrapopulation variation of both morphological traits and shell banding polymorphism using the example of a continuous population (a metapopulation) of the land snail Cepaea vindobonensis (Férussac, 1821). A total of 14 samples of the land snail C. vindobonensis from a population located in the “Dubki” Park (Ukraine, Mykolayiv) were collected in May-July 2007. The land snail collection sites are divided into three areas by buildings and asphalt roads, indicated by the Latin letters A, B and C. The major diameter of shell (MJD), the minor diameter of shell (MID) and the shell height (SH) were measured with a digital calliper to the nearest 0.05 mm. Two shell shape indices (SF1 and SF2) were also computed. A high level of the intrapopulation variation of the land snail C. vindobonensis was found in our study. Significant differences between sample means were found for all shell traits and indices used (except for SF1), however, maximum difference was noted for SH and SF2 (in both cases: P ≤ 0.001). About 80% of the total variation of the variance-covariance matrix was explained by the 1st and 2nd Principal Components (PC1 and PC2). The PC1 was characterized by high positive factor loadings of MJD, MID and SH and thus can be interpreted as “shell size dimension” and the PC2 had a high correlation with SF2 and thus it can be interpreted as “shell globularity”. The PC1 and PC2 determined a high level of spatial differentiation of intrapopulation morphological variation of the land snail C. vindobonensis. Samples with small (area A) and large (areas B and C) shells were separated from each other according to the PC1. Areas B and C were characterized by individuals with flatness and globularity shells, respectively. The areas A, B and C differed significantly in the total number of morphs, average number of morphs and frequency of rare morphs (Kruskal-Wallis H-test; in all cases P ≤ 0.010). At the same time, the highest value of phenetic diversity was noted for samples collected within A and C areas. In general all phenetic diversity estimators showed a positive correlation with sample size (Spearman's correlation coefficient; in all cases P ≤ 0.05). It was found that the type of biotope did not likely affect the frequency of individual morphs with respect to the shell banding polymorphism pattern. With regard to the most common morphs two patterns of the spatial arrangement of the intrapopulation variation were found – clinal pattern was for the frequency of pallescens morph and chaotic pattern was for frequency of “12345” morph.

Snail shell colour evolution in urban heat islands detected via citizen science

The extreme environmental conditions that prevail in cities are known to cause selection pressures leading to adaptive changes in wild, city-dwelling, organisms (urban evolution). The urban heat island, elevated temperatures in the city centre due to a combination of generation, reflection, and trapping of heat, is one of the best recognised and most widespread urban environmental factors. Here, we use a citizen-science approach to study the effects of urban heat on genetically-determined shell colour in the land snail Cepaea nemoralis in the Netherlands. We use smartphone applications to obtain colour data on almost 8000 snails throughout the country. Our analysis shows that snails in urban centres are more likely to be yellow than pink, an effect predicted on the basis of thermal selection. Urban yellow snails are also more likely to carry dark bands at the underside of the shell; these bands might affect thermoregulation in yet underexplored ways.

Discrete or indiscrete? Redefining the colour polymorphism of the land snail Cepaea nemoralis

Biologists have long tried to describe and name the different phenotypes that make up the shell polymorphism of the land snail Cepaea nemoralis. Traditionally, the view is that the ground colour of the shell is one of a few major colour classes, either yellow, pink or brown, but in practise it is frequently difficult to distinguish the colours, and define different shades of the same colour. To understand whether colour variation is in reality continuous, and to investigate how the variation may be perceived by an avian predator, we applied psychophysical models of colour vision to shell reflectance measures. We found that both achromatic and chromatic variation are indiscrete in Cepaea nemoralis, being continuously distributed over many perceptual units. Nonetheless, clustering analysis based on the density of the distribution did reveal three groups, roughly corresponding to human-perceived yellow, pink and brown shells. We also found large-scale geographic variation in the frequency of these groups across Europe, and some covariance between shell colour and banding patterns. Although further studies are necessary, the observation of continuous variation in colour is intriguing because the traditional theory is that the underlying supergene that determines colour has evolved to prevent phenotypes from “dissolving” into continuous trait distributions. The findings thus have significance for understanding the Cepaea polymorphism, and the nature of the selection that acts upon it, as well as more generally highlighting the need to measure colour objectively in other systems.

Recombination within the Cepaea nemoralis supergene is confounded by incomplete penetrance and epistasis

Although the land snail Cepaea nemoralis is one of the most thoroughly investigated colour polymorphic species, there have been few recent studies on the inheritance of the shell traits. Previously, it has been shown that the shell polymorphism is controlled by a series of nine or more loci, of which five make a single ‘supergene’ containing tightly linked colour and banding loci and more loosely linked pigmentation, spread band and punctate loci. However, one limitation of earlier work was that putative instances of recombination between loci within the supergene were not easily verified. We therefore generated a new set of C. nemoralis crosses that segregate for colour, banding and pigmentation, and several other unlinked shell phenotype loci. The snails were genotyped using a set of RAD-seq-derived loci that flank the supergene, and instances of recombination tested by comparing inferred supergene genotype against RAD-marker genotype. We found no evidence that suspected ‘recombinant’ individuals are recombinant between loci within the supergene. As point estimates of recombination between both colour/banding, and colour/pigmentation loci are zero, incomplete penetrance and epistasis are a better explanation for the apparent ‘recombinant’ phenotype of some snail shells. Overall, this work, therefore, shows that the architecture of the supergene may not be as previously supposed. It also provides a resource for fine mapping of the supergene and other major shell phenotype loci.

Intraspecific morphological variation of the sperm storing organ in two hermaphroditic land snail species

BackgroundPostcopulatory sexual selection is very important in species with reproductive strategies that involve multiple mating and prolonged sperm storage. The sperm storage organ has been hypothesized to evolve in response to different levels of sperm competition in several species while population density has been considered as a factor that approximates sperm competition risk and intensity in the field. Apart from population density, local microclimatic conditions may also play a role in determining sperm competition levels in natural populations of land snails by affecting their chances of encountering mates. The goal of this study was to investigate the variation of the structure of the sperm storage organ in the simultaneously hermaphroditic land snails Helix lucorum and Cepaea vindobonensis occurring sympatrically in two sites which differed in habitat humidity. The populations of both species from the two sites, also differed in density and in duration of reproductive period. Multiple samples were taken from each population in order to test for temporal variation.ResultsIn both species, the spermatheca consisted of a simple fertilization chamber and a variable number of lateral tubules. The length of the spermatheca showed no temporal or spatial differentiation nor did it show any correlation with snail size. The number of tubules in Helix lucorum ranged from five to sixteen and in Cepaea vindobonensis from one to eight and in both species a significant difference of this trait between the two study sites was detected. In Cepaea vindobonensis, the difference in tubule number led to difference of the total tubule length which reflects sperm storage capacity of the spermatheca but this was not the case with H. lucorum in which no increase in total tubule length was detected.ConclusionsIntraspecific variation in the spermatheca was observed in both snail species studied. The variation observed was independent of snail size, and reproduction status, while the two species responded differently to higher sperm competition levels.

A century after introduction: variability in Cepaea hortensis (Müller, 1774) in Sibiu, central Romania

AbstractThe polymorphic land snail Cepaea hortensis was introduced to the city of Sibiu, central Romania, in the first decade of the 20th century and has spread widely across the city. A total of 97 locations were examined in 2017 across the city to determine the habitat preferences and variation in shell size, shape, colour and banding polymorphism of C. hortensis, and to relate these to the same features in the likely source population from Mannheim, Germany, and the first established population in Sibiu. We found that C. hortensis was largely restricted to sites with some woody vegetation cover and showed a marked preference for abandoned and overgrown private gardens. Mean adult shell size in present-day populations was almost always smaller than in both the presumed population of origin from Germany and the first recorded population from Sibiu. Populations showed a wide range of variation in frequencies of shell colours and banding morphs. This variation was not related to habitat or cover, and there was no evident geographical structure in the patterns. Comparisons with a smaller-scale sampling in 2004/5 showed that some populations had gone extinct, some remained stable and in some morph frequencies had changed drastically, but in no consistent direction. These results are similar to those obtained for the related species C. nemoralis in comparable circumstances, but differ from those obtained from regions where C. hortensis is long established and where habitats have been stable. The patterns we observed most likely reflect the effect of passive dispersal by humans, genetic drift and founder effects.

Shell colour, temperature, (micro)habitat structure and predator pressure affect the behaviour of Cepaea nemoralis

Although shell colour polymorphism of the land snail Cepaea nemoralis is a well-known phenomenon, proximate and ultimate factors driving its evolution remain uncertain. Polymorphic species show variation in behavioural responses to selective forces. Therefore, we estimated effects of various environmental factors (temperature, humidity, food availability, (micro)habitat structure and predatory pressure) on behavioural response (frequency of locomotion, climbing and hiding) of C. nemoralis morphs, in experimental and natural conditions. In the experimental part of study, the frequency of locomotion was negatively affected by temperature and the presence of food and positively influenced by the presence of light. Morphs significantly differed in behavioural responses to environmental variability. Pink mid-banded and yellow five-banded morphs climbed less often and hide in shelter more often than yellow and pink unbanded individuals when temperature was low and food was absent. Snails fed most often at moderate temperature compared to low and high temperatures. Field investigations partially confirmed differences among morphs in frequency of climbing, but not in terms of probability of hiding in sheltered sites. In natural colonies, temperature and (micro)habitat structure significantly affected frequency of climbing as well as hiding in shelter. Snails more often hid in sheltered sites where thrushes preyed on Cepaea. Tendency of unbanded morphs to climb trees may have evolved under avian predatory pressure as thrushes forage on a ground. Tendency of banded morphs to hide in sheltered sites may reflect prey preferences for cryptic background. The results implicate that differential behaviour of C. nemoralis morphs compensate for their morphological and physiological limitations of adaptation to habitat.

Inferring microevolution from museum collections and resampling: lessons learned from Cepaea.

Natural history collections are an important and largely untapped source of long-term data on evolutionary changes in wild populations. Here, we utilize three large geo-referenced sets of samples of the common European land-snail Cepaea nemoralis stored in the collection of Naturalis Biodiversity Center in Leiden, the Netherlands. Resampling of these populations allowed us to gain insight into changes occurring over 95, 69, and 50 years. Cepaea nemoralis is polymorphic for the colour and banding of the shell; the mode of inheritance of these patterns is known, and the polymorphism is under both thermal and predatory selection. At two sites the general direction of changes was towards lighter shells (yellow and less heavily banded), which is consistent with predictions based on on-going climatic change. At one site no directional changes were detected. At all sites there were significant shifts in morph frequencies between years, and our study contributes to the recognition that short-term changes in the states of populations often exceed long-term trends. Our interpretation was limited by the few time points available in the studied collections. We therefore stress the need for natural history collections to routinely collect large samples of common species, to allow much more reliable hind-casting of evolutionary responses to environmental change.

Candidate genes for shell colour polymorphism in Cepaea nemoralis.

The characteristic ground colour and banding patterns on shells of the land snail Cepaea nemoralis form a classic study system for genetics and adaptation as it varies widely between individuals. We use RNAseq analysis to identify candidate genes underlying this polymorphism. We sequenced cDNA from the foot and the mantle (the shell-producing tissue) of four individuals of two phenotypes and produced a de novo transcriptome of 147,397 contigs. Differential expression analysis identified a set of 1,961 transcripts that were upregulated in mantle tissue. Sequence variant analysis resulted in a set of 2,592 transcripts with single nucleotide polymorphisms (SNPs) that differed consistently between the phenotypes. Inspection of the overlap between the differential expression analysis and SNP analysis yielded a set of 197 candidate transcripts, of which 38 were annotated. Four of these transcripts are thought to be involved in production of the shell’s nacreous layer. Comparison with morph-associated Restriction-site Associated DNA (RAD)-tags from a published study yielded eight transcripts that were annotated as metallothionein, a protein that is thought to inhibit the production of melanin in melanocytes. These results thus provide an excellent starting point for the elucidation of the genetic regulation of the Cepaea nemoralis shell colour polymorphism.

The Efficiency of Flotation Compared with Other Methods for Recovering Assemblages of Terrestrial and Aquatic Gastropods from Archaeological Deposits, with Reference to the Site of Pico Ramos (Basque Country, Spain)

ABSTRACTWe analysed assemblages of shells of land snails and small aquatic and wetland gastropods from a late Mesolithic-early Neolithic deposit in the cave of Pico Ramos (Basque Country). Shells were recovered by hand during excavation and all excavated deposit was processed by flotation, with a 250 μm mesh to recover floating material and a 1.0 mm mesh to recover the heavy fraction. The total assemblage comprised 5780 individuals (MNI) among 33 taxa, the sieved sub-assemblage having the highest proportions of these (2841 MNI in 31 taxa) and the hand-picked sub-assemblage the lowest (698 MNI in 11 taxa). Eleven taxa were absent from the flotation sub-assemblage, although the recovered MNI (2241; 39% of the total MNI) was high. The palaeoecological implications of the results are considered. Adult and large-sized juvenile shells of the edible land snail Cepaea nemoralis are abundant in the hand-picked sub-assemblage, but it is the absence of small-sized juvenile shells in the flotation and wet-sieved sub-assemblages that permits the inference that the species was collected for consumption. Wet sieving is therefore essential, both for interpreting hand-picked samples and in particular for the recovery of representative assemblages of land molluscs.

Land snails benefit from human alterations in rural landscapes and habitats

AbstractMetapopulation functioning in the context of landscape and habitat structure is generally well understood. However, in human‐transformed landscapes, human disturbance may affect not only a species of interest but also biotic interactions in which that species is engaged, which has been rarely studied at multiple spatial scales. We tested (1) the associations between local (habitat and micro‐habitat) and landscape factors and the occurrence and abundance of the land snail Cepaea nemoralis and two of its predators (thrushes and mice); and (2) the hypothesis that human disturbances modify associations between C. nemoralis and its predators. We surveyed snail colonies in rural landscapes of Poland. Generalized linear mixed models were used to test the stated hypotheses. At the landscape scale, the occurrence of C. nemoralis colonies was positively related to the cover of suitable habitats, fallow/waste land, and road length in the surrounding landscape. Bird predator abundance was positively correlated with the occurrence of snail colonies, mostly at sites with a high human impact, and was negatively correlated with snail abundance at sites with low human disturbance. Snail occurrence and abundance at the micro‐habitat scale were positively correlated with variables describing the proximity of shelter and humidity. Avian predation pressure was negatively correlated with forest and settlement cover, whereas rodent predation pressure was positively associated with snail abundance and was negatively correlated with bird abundance. Human disturbance at the habitat and landscape scales benefited C. nemoralis through the creation of suitable habitats and scaling down of avian predation pressure. Conditions related to shelter and humidity were linked to the distribution and abundance of snails at the micro‐habitat scale.

Differential shell strength of Cepaea nemoralis colour morphs--implications for their anti-predator defence.

One of the most spectacular evolutionary forces is predation, evidenced to stimulate polymorphism in many prey species. Shell colour polymorphism of the land snail Cepaea nemoralis is a well-known model in evolutionary research. Nevertheless, the knowledge on the ecological causes driving its evolution remains incomplete and proximal factors shaping predatory pressure on C. nemoralis morphs are unknown. We evaluated shell crushing resistance and thickness, constituting crucial snail anti-predator defences in two shell areas (the apex and labium) of eight C. nemoralis morphotypes differing in shell colour and banding pattern. A GLM showed a significant effect of shell colour, banding pattern and shell thickness on shell strength. Pink shells were stronger than yellow ones, and banded forms had stronger shells than unbanded snails. The labium (usually attacked by mice) was generally thicker and more resistant than the apex (usually crushed by birds). Thicker shells were more resistant to crushing, and the rate of shell strength increase per unit of shell thickness was greater in pink and banded individuals compared to yellow and unbanded ones. Yellow and unbanded morphs have been found to be preferred by mice in the previous studies, which suggests that shell strength may be an important trait used in prey selection by these shell-crushing predators. The differences in potential anti-predator defences among snail morphs, found in the present study, justify future research on direct effect of C. nemoralis morphs shell strength on predator selectivity.