Self-fertilization and cross-fertilization in the land snail Arianta arbustorum (Mollusca, Pulmonata: Helicidae)

The simultaneously hermaphroditic land snail Arianta arbustorum (L.) (Pulmonata: Helicidae) was long believed to be self-incompatible. In the present study, however, uniparental reproduction was observed under laboratory conditions. Snails from two natural populations and from laboratory culture reared in isolation from the subadult stage produced fertile eggs. Over a period of three years after maturation, 38.5% of the unmated snails produced fertile eggs, mostly in the second and third years instead of the first year. Egg production and hatching success (the proportion of eggs that hatched) were lower in unmated snails than in mated snails. The number of hatchlings produced by unmated snails was 1–2% of that produced by mated snails. Hatchlings from unmated snails survived as well as those from mated snails under laboratory conditions. The result indicate that A. arbustorum can self-fertilize but with a great fitness reduction.     

Proximate factors influencing egg cannibalism in the land snail Arianta arbustorum (Pulmonata,Helicidae)

Summary Byttneria aculeata (Sterculiaceae), a subcanopy liane with a shrubby juvenile form has two distinct leaf color morphs in juvenile plants- a given juvenile has plain green leaves or leaves with whitish variegation. Both forms occur together in the forest and in clearings; however, the variegated morph is more common in open sites, and the plain morph predominates in the forest. Percent variegation per leaf for variegated plants increased from closed to open sites. Measurements of growth support the idea that variegation is favored in open habitats. Within a given habitat, rate of herbivory by leaf miners on a given morph increases with increasing relative frequency of that morph; however, at a given relative frequency, the variegated morph is less heavily attacked than is the plain morph when it occurs elsewhere at that same frequency.     

Induced environmental variation in Arianta arbustorum (L.)

By raising Arianta from egg to adult under 4 different conditions environmental variation was produced. Stocks used were from midland and alpine places in Switzerland. The study involves 6 measurements and 6 coded characters. In spite of all variation, which was consistent in some characters, the stocks maintained their identity.     

Thermal advantage of pale coloured morphs of the snail Arianta arbustorum (Helicidae, Pulmonata) in alpine habitats

In alpine grasslands pale colour morphs of the snail Arianta arbustorum are more frequent than dark ones As pale morphs absorb less solar radiation, they may forage for longer in sunlight and/or have a lower mortality risk from overheating This hypothesis was tested Equal numbers of pale and dark morphs were kept in enclosures in an alpine meadow During the day, more pale morphs were seen than dark ones, the latter obviously remaining hidden The difference was not significant in enclosures set up in a nearby forest, confirming that the inequality in behaviour depends on the amount of sunlight Dark snails are on average warmer than pale ones
Experiments were also conducted in grassland on a mountain slope at altitudes between 1200 and 2000 m Snails were marked with a paint which fades in light, and released When recaptured several days later, the marks were more faded in pale morphs than in dark, suggesting that they had been more exposed to sunlight, supporting our hypothesis The difference in fading was more pronounced at high than at low altitude     

The Pleistocene glaciations and the evolutionary history of the polytypic snail species Arianta arbustorum (Gastropoda, Pulmonata, Helicidae)

The evolutionary history of the snail Arianta arbustorum is controversial. This diverse, polytypic species has two distinct forms: one, with a globular shell and closed umbilicus, is found from lowland to high altitudes; the other, with a depressed shell and open umbilicus, is found at a few scattered, high altitude localities. What is the origin of these two forms? Some believe that the depressed shell is a recent, local, ecotypic adaptation to alpine environments. Others believe that this form is a relic of an ancestral condition that may have survived the Pleistocene glaciations on nunatak-like montane refugia, while the globular shell is a derived condition and its presence at high altitudes follows post-Pleistocene recolonisation. We analysed a portion of the mitochondrial gene cytochrome oxidase I for 100 snails of the species A. arbustorum, three additional Arianta species, and nine outgroup taxa from five genera, in order to understand the phylogeographic history of the species. Despite some confounding artefacts that are likely due to introgression among the morphological forms, the resulting phylogeny shows that the depressed shell is plesiomorphic, while the globular shell is derived. Moreover, their disparate histories suggest that the depressed shell variety survived the glaciations in pockets of alpine refugia, while the globular shell variety recolonised the alpine environment post-glacially.     

The spermatheca in the land snail, Arianta arbustorum (Pulmonata: Stylommatophora): muscle system and potential role in sexual selection

Abstract. Previous studies have shown variable patterns of paternity after multiple mating, and also variation in sperm storage among individuals of Arianta arbustorum , which suggests that the spermatheca may influence paternity in this promiscuous land snail. To identify possible morphological correlates of sperm manipulation, we investigated arrangement and ultrastructure of the muscles of the spermatheca. The musculature surrounding the 2–9 spermathecal tubules is arranged in a complex three dimensional network. In addition, each tubule has a thin sheath in which longitudinally oriented cells make up the innermost layer. Usually, the smooth muscle cells are enclosed by connective tissue. Only occasionally is direct muscle-muscle contact established through dense plaques. The short thick filaments, their small diameter, the relatively weak development of the tubular system and sarcoplasmic reticulum, and the low density of mitochondria indicate that the muscle cells contract relatively fast but with little strength, that they recover slowly, and have low endurance. A single muscle cell may be innervated by several axons and one axon may contact several muscle cells. Combining evidence of the present paper and a foregoing investigation on the spermathecal epithelium, we suggest that the main function of the spermathecal muscles is to expel sperm stored for fertilization, while the ciliation of the common duct is probably responsible for the distribution of sperm among the tubules.     

Variation of distal genitalia in the simultaneously hermaphroditic land snail Arianta arbustorum (Pulmonata,Stylommatophora) caused by sexual selection?

The most likely explanation for genitalic extravagance may be sexual selection acting either before, during, or after copulation. In promiscuous species, the intensity of postulating sexual selection may be a function of population density. In this study we examined the variability of the distal genitalia involved in spcrmatophore production, reception and manipulation of 113 adult individuals of the simultaneously hermaphroditic land snail Arianta arbustorum (L.) from six natural populations in the Eastern Alps (Gesause, Austria). We investigated the hypothesis that these genitalia increase in relation to shell size with increasing population density (range: 0.9 to 39.8 individuals/m²) and expected a higher variance of the genitalia compared to shell dimensions due to sexual selection. Genitalic size was unexpectedly inversely related to population density, probably due to increased inhibitor)^- effects of snail mucus. Patterns of variation of female and male characters did not differ. Coefficients of variation of the genitalia were significantly higher than those of the shell dimensions as predicted. This was due to a higher dispersion around the regression lines rather than higher allometric values. However, the influence of sexual selection on genitalic size and variance cannot be unambiguously determined. We discuss different scenarios emphasizing the importance of sexual selection to differing degrees and conclude that sexual selection has probably played only a minor role.     

Altitudinal distribution of alien plant species in the Swiss Alps

In summer 2003 we recorded the presence and abundance of alien plant species at 232 sites (107 railway stations and 125 road sites) along mountain passes in the Swiss Alps. The altitudinal distribution of species was related to the current abundance of the species in Switzerland and time since introduction. A total of 155 alien taxa were recorded. Numbers of species per site declined exponentially with altitude, and only a few species were found in the alpine zone (>2000 m). In contrast, species richness among comparable native taxa appeared to be nearly independent of altitude over the range investigated. Maximum altitude reached by alien species was related positively to both total area occupied in Switzerland and to time since introduction. A comparison of the results with earlier records suggests that many species, particularly those previously restricted to low or intermediate altitudes, have advanced their altitudinal limits over the past few decades. Various hypotheses are presented to explain the declining abundance of alien species with altitude: low-altitude filter effects, low propagule pressure, and genetic swamping of peripheral populations at higher altitudes. However, at present we do not have sufficient evidence to determine the relative importance of these effects. We conclude that invasion into mountain areas such as the Swiss Alps tends to proceed rather slowly, though the process may be accelerated by climatic warming. For this reason, further research to investigate the processes determining how plants invade mountain areas is urgently needed. And more generally, investigations into the distribution of alien species along strong altitudinal gradients may provide valuable insights into the mechanisms driving the spread of alien organisms.     

Allozymic differentiation in the land snail Arianta arbustorum (Stylommatophora, Helicidae): historical inferences

The intraspecific genetic differentiation depends to a large part on historical factors. The mode of dispersal is an important determinant of the genetic diversity especially of newly founded populations. Fast long-distance colonization of formerly unsuitable habitats results in reduced genetic diversity and lineages originating from different glacial refugia form well-defined hybrid zones, where they meet. Slow dispersers, on the other hand, preserve their refugial genetic diversity. Paleontological findings and a previous analysis of mtDNA suggest that the land snail Arianta arbustorum, which today is widely spread across Europe, has survived the Pleistocene glaciations within the boundaries of permafrost. In the present paper, we asked, whether the differentiation of nuclear markers, i.e. allozymes, supports these results. We investigated the allozymic differentiation at 15 polymorphic loci in 14 Alpine and one English populations of the land snail A. arbustorum. Various tests of genetic population differentiation yielded ambiguous results. Allele frequencies differed significantly across populations and all but five pairwise F_ST values were significant. Genetic distances between populations were comparatively high. On the other hand, an exact test of population differentiation found no differences and the assignment of individuals to their populations of origin was poor. This indicates that allozymic differentiation is constrained. Whether these constraints are of natural or methodological nature remains open. Matrices of genetic and geographical distances were correlated only after removal of the English population from the analysis. Similarly, with one exception F_ST values increased as the number of regions included in the calculations increased. Thus, on a regional scale, we largely observed a pattern of isolation by distance, which does no longer hold on a larger scale. This is well in accordance with the previous findings based on mtDNA. Mitochondrial lineages were widely distributed across Europe and overlapped extensively. Considering the low potential for dispersal of a land snail, the patchy distribution of lineages, and the high genetic diversity also found in the present study suggests that A. arbustorum has survived the Pleistocene within the boundaries of permafrost.     

Interpopulation variation in the prevalence and intensity of parasitic mite infection in the land snail Arianta arbustorum

Abstract. The parasitic mite Riccardoella limacum sucks blood in the lung of its host, the land snail Arianta arbustorum . The infection of various host populations was examined in Switzerland. In a lowland snail population, prevalence of infection did not vary among seasons. However, intensity of mite infection in dissected individuals of A. arbustorum was high in autumn, but low in winter and spring when ≤100 mite eggs were found attached to the lung epithelium. A novel, non-invasive parasite screening method was used to estimate the number of mites on living host snails. An analysis of repeatability revealed that 92.9% of the snails were correctly classified as infected or non-infected with this non-invasive method. Prevalence of mite infection was examined in 997 adults of A. arbustorum from 11 natural populations distributed over an altitudinal gradient ranging 335–2360 m. No infected snails were found in 7 populations, while in the remaining 4 populations the prevalence of mite infection ranged 45.8–77.8%. Intensity of infection also differed among the 4 host populations. No geographic pattern in prevalence of infection was found. However, parasitic mites did not occur in snail populations situated at elevations of 1290 m or higher. A possible explanation for this finding could be that the host's hibernation period may be too long at high elevations for mites and their eggs to survive. At low elevations, other factors may affect the presence of R. limacum in the lung of A. arbustorum .     

Ultrastructure of the osphradium of Siphonaria grisea (Mollusca, Pulmonata)

By means of scanning and transmissive electron microscopic methods osphradium of Siphonaria grisea has been studied. The osphradium of the animal is presented as a small torulus formed by supporting ciliated cells. Among them bodies of receptory cells are situated; they are of smaller size and decorated with a bundle of microvilli. Central processes of the receptory cells, penetrating through a thick layer of the connective tissue, reach the subepithelial neural trunk. Concentration of the neural cells in the periphery of the trunk is small, and in the central area no chemical synapses are revealed. The data presented demonstrate a primitive structure of the osphradial chemoreceptory organ in Siphonaria.     

Epiphallus anatomy in the Arion hortensis species aggregate (Mollusca, Pulmonata)

A SEM study of the structure associated with the outlet of the epiphallus in the atrium genitale of three arionid species ( Arion hortensis s.s., A. distinctus and A. owenii ) is presented. The structures are described and compared, and the specific differences are outlined. Preliminary notes on the same structure in A. intermedius , as well as on the spermatophores of A. hortensis s.s. and A. distinctus , are also provided. It is concluded that A. owenii stands somewhat apart in relation to the latter two species, since it may be more related to A. intermedius. Although it is generally accepted that the genus Arion is characterized by the absence of a penis, it is suggested here that the described structure at the outlet of the epiphallus is nothing but a remnant of an ancestral penis; anyhow, it is at least functionally allied to a penis.     

Patterns of dispersion, density and dispersal in alpine populations of the land snail Arianta arbustorum (L.) (Helicidae)

In studying populations of the land snail Arianta arbustorum in the eastern Swiss Alps, juvenile and subadult snails were found to be aggregated all the time, while adults were mostly aggregated during summer, the main mating period. The mean size of aggregations was 25 cm × 25 cm but varied, according to the area a grass tuft covered, while the intensity of aggregations was influenced by weather conditions. Densities of 10–15 subadults and 3–6 adults m⁻² were found along ditches and streams, 2–25 subadults and 2–20 adults on uncultivated alpine meadows, while on scree-covered alpine grassland the density of subadults and adults was 0.3–0.5 m⁻². The extent of daily movements varied with microclimatic factors and season and depended on the structure of the habitat. The dispersal rate was greatest in June (2.6 m month⁻¹) and least in August (1.1 m month⁻¹). In summer and autumn, active uphill movements were noted, a process which compensates for the loss of altitude by passive downhill displacements. Roiling down on snowfields and being carried away by avalanches and streams were observed.     

Biochemical variation and shell shape in populations of the fresh-water snail Lymnaea peregra (Mollusca, Gastropoda, Pulmonata) from south-west Ireland

Lymnaea peregra was collected at sites, predominantly from the Caha plateau, in south-west Ireland. These included sites investigated by previous workers and found to contain populations with a distinctive pattern of variation in shell-shape. Normally spired populations and non-spired ( involuta, sub-involuta ) could occur in adjacent loughs in the same watershed. The biochemical variation in some of these populations was investigated using horizontal starch-gel zone electrophoresis. Fourteen biochemical loci were investigated. Although circumscribed by difficulties in collecting and low sample numbers, the investigation showed that the pattern of variation in shell shape was still present and was not correlated or associated with the biochemical variation. While this did not support hypotheses of geographic race formation or incipient speciation, it was in agreement with the current view of variation and speciation in fresh-water pulmonates.     

Expandable spermatheca influences sperm storage in the simultaneously hermaphroditic snail Arianta arbustorum

Summary

In many simultaneously hermaphroditic land snail species, the sperm storage organ (spermatheca) is highly structured, suggesting that the female function might be able to influence offspring paternity. Physical properties of the sperm storage organ, including its initial size and sperm storage capacity, may also affect fertilization patterns in multiply mated snails. We examined the structure, volume and tubule length of empty spermathecae in the land snail, Arianta arbustorum, and assessed differences in spermatheca size following a single copulation. The number of spermathecal tubules ranged from 2–7, but was not correlated with the volume of empty spermathecae. The volume of sperm stored in the spermatheca after a copulation was correlated with neither the number of spermathecal tubules nor copulation duration. Mean spermathecal volume more than doubled between two and thirty-six hours after sperm uptake, but the length of the spermathecal tubules did not change. Interestingly, the volume of sperm stored in the spermatheca seems not to be related to the size of the spermatophore and thus not to the number of sperm received (= allosperm). The amount of allosperm digested in the bursa copulatrix was highly variable and no significant relationship with the size of the spermatophore received was found. These findings suggest that numerical aspects of sperm transfer are less important in influencing fertilization success of sperm in A. arbustorum than properties of the female reproductive tract of the sperm receiver.