Shell thickness in the land snail Theba pisana (Pulmonata: Helicidae)

Shell thickness in Theba pisana was measured in samples from Britain, northern France and the Mediterranean region. Mean adult thickness varied between about 0.08 mm and 0.20 mm. These values are much lower than previously published values. The considerable variation within samples was related to shell size, periods of growth, and resorption of shell material during breeding. Variation between samples was not great. This contrasts with previous work which linked shell thickness variation to large-scale climatic effects. Perhaps availability of calcium may be more important than climate in determining shell thickness in Theba Pisana .     

Thirty-four years of climatic selection in the land snail Theba pisana

Few continuous, long-term studies have measured the intensity and variability of natural selection within a framework of clear adaptive hypotheses. In the snail Theba pisana, the proportion of effectively unbanded shells is higher in exposed habitats than in adjacent acacia thickets, which has been explained by microclimatic selection. Comparisons across an ecotone for 34 consecutive years determined the combined effects on morph frequencies of habitat and changes in weather conditions during summer. The long-term average (±s.e.) frequency of effectively unbanded shells was 0.577±0.011 in the open habitat when compared with 0.353±0.005 in the acacia. The persistent association of shell banding with habitat accounted for 34% of the variation in morph frequencies. Differences among years were also large, representing 23% of the variation. Higher proportions of effectively unbanded snails were associated with hotter, sunnier summers. Thus, temporal variation supports the hypothesis of microclimatic selection, consistent with the spatial association with habitat. Based on observed rates of change, the mean annual selection on this polymorphism was about 0.13, but with a large variance: s was as high as 0.5, but ⩽0.05 in about 40% of the years. The large variance and frequent reversals in direction of selection indicate a potential for rapid genetic change, but with little net change in morph frequencies over three decades, highlighting the value of long-term continuous studies of populations facing natural environmental variation.     

Epistasis, phenotypic disequilibrium and contrasting associations with climate in the land snail Theba pisana

Hotter conditions favour effectively unbanded (EUB) shells in the snail Theba pisana. T. pisana is also polymorphic for colour of the shell's apex, determined by a pair of alleles at a locus linked to the banding locus. Apex colour is epistatic to shell banding, such that banded snails with a dark apex have darker bands. Annual censuses over 22 years across an ecotone between a sheltered Acacia thicket and open dune vegetation showed a persistent association of both EUB shells and pale apex with the Open habitat. The parallel variation was due partly to strong phenotypic disequilibrium, as the combination of EUB with dark apex was rare. Nevertheless, in fully banded shells the frequency of pale apex was also higher in the Open habitat, confirming independent, parallel associations of the two contributors to paleness. Within the Acacia habitat, temporal variation of the frequencies of banding morphs was much greater than for apex colour, and EUB shells were associated with hotter summers. Consistent with its primary effect only on the very small snails, apex colour did not vary with summer conditions, but instead, higher frequencies of pale apices were associated with sunnier winters. The intensity of selection was lower on apex colour than shell banding, due partly to the constraint of phenotypic disequilibrium. The shell traits in T. pisana are an example of complex responses to climatic variation, in which phenotypic disequilibrium constrains evolution of apex colour, but separate mechanisms of selection are evident.     

Regulation of infection by a nematode (Muellerius capillaris) in the snail Theba pisana

The regulation of infection by Muellerias capillaris was studied in three groups of Theba pisana. The regulation prior to penetration was very important for all the groups as assessed by the use of snail-conditioned water: a lytic activity was observed and the surviving larvae were less active than the control. The mortality rate of larvae after penetration was low in preinfected local group of snails, medium in the naive nonlocal group, and high in the naive local group.     

Historic data analysis reveals ambient temperature as a source of phenotypic variation in populations of the land snail Theba pisana

Pulmonate land snails are often polymorphic in their shell coloration pattern. To quantify the contribution of environmental parameters to the nondirectional change in phenotypic variation, we used a historic dataset on Theba pisana morph frequencies and climate data for statistical modelling. We found significant correlations of the degree of phenotypic diversity between juveniles and corresponding adult individuals within the same and the subsequent generation. Among climate parameters, the phenotypic diversity of adults correlated significantly and positively with the mean and maximum ambient temperatures in the winter and spring only. There was no correlation between high or low temperatures and the frequency of distinct morphs. Akaike's information criterion‐based model selection revealed the particular importance of only parental phenotypic diversity for next generation juvenile phenotypic diversity. By contrast, phenotypic diversity of the juveniles of the preceding year and the mean temperatures in winter and spring were important for the phenotypic diversity of adult snails. Approximately two‐thirds of the explicable variation in phenotypic diversity of adults was explained by inheritance and approximately one‐third was expained by ambient temperature. The present study shows that genetics and temperature interact to generate nondirectional changes in phenotypic variation within populations, which also can be reflected by changes in the phenotype of individuals. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 109 , 241–256.     

Yeasts and fungi in excrements of the sand hill snail,Theba pisana

Eight microbial species were isolated and identified from excrements of the sand hill snail,Theba pisana, viz. 3 filamentous fungi, 3 basidiomycetous yeasts and 2 yeast-like ascomycetous fungi.     

Natural history of Theba pisana in Israel (Pulmonata: Helicidae)

Theba pisana in southern Israel is a semelparous species with an annual life cycle. Young snails hatch in early winter, grow rapidly in spring and by summer they reach, first adult shell-size, then adult shell-weight. Shell-size is negatively related to density, suggesting that, as in some other landsnails, there may be a density-dependent inhibition of growth. Summer forces the snails into 3–4 months of aestivation.
After the first winter rains the snails, awakened perhaps by the sudden drop in barometric pressure, emerge from aestivation. Only now are most of the spermatozoa and mature oocytes formed (maturation of gametes actually starts some two weeks before the first rains), and the snails copulate. No evidence of precocious breeding was found. The snails sometimes dig a few "sham" holes, perhaps to discourage egg-eating predators, before finally depositing their eggs (mean clutch size: 80 eggs) in the last hole.
In mid-winter, the snails die. In a habitat where the sheltering conditions of the vegetation offer the snails more protection, the dying off process of the population is stretched over a few more months. This could, perhaps, be the first step towards the evolution of a bi-annual cycle, such as is typical of T. pisana in some other areas of its distribution.     

The effect of temperature on the activity of three species of Helicid snail (Mollusca: Gastropoda)

The activity of the land snails Cepaea nemoralis (L.), C. hortensis (Müll.) and Arianta arbustorum (L.) was measured over 24 hours at the constant temperatures of 0, 3, 8, 17, 22 and 26°C at 100% relative humidity and with 16 hours light and 8 hours dark. The activity of all three species varies significantly with temperature and light, and there are differences in the activities of the species at each temperature. A. arbustorum is relatively more active than Cepaea at low rather than high temperatures, as is C. hortensis with respect to C. nemoralis , though to a lesser extent. There is little diurnal rhythm in activity at 0°C, but all species become more nocturnal as temperature increases; C. nemoralis is most, and A. arbustorum least nocturnal. The results obtained assist in explaining the observed geographical and local distributions of the species concerned.     

New insights into and limitations of the molecular phylogeny in the taxon-rich land snail genus Montenegrina (Mollusca: Gastropoda: Clausiliidae)

Rock-dwelling gastropods are usually patchily distributed in limestone habitats, presumably have low active and passive dispersal ability and often represent narrow-ranged endemic taxa. Their current taxonomy is predominantly shell morphology based, and it remains unknown whether the morphologically differentiated and geographically separated populations represent phylogenetic clades. In this study, we analysed the hyperdiverse, terrestrial door snail genus Montenegrina. Based on the current taxonomy defined by shell morphology, it contains 29 species and 106 subspecies distributed in the Balkan region. The constructed phylogenetic tree using three mitochondrial markers was used to test whether it agrees with the current taxonomy. In this comprehensive tree, about half of the species and subspecies are monophyletic. Some of the paraphylies could be reasonably resolved by taxonomic changes; that is, some subspecies should be reassigned or raised to species level. Other incongruencies probably arose due to introgression even between distant clades. The histone genes turned out to be unsuitable for elucidating the phylogeny of Montenegrina. In the species-delimitation tests, considerably more molecular operational taxonomic units were delimited than the number of presently described species. The present data indicate that (a) shell morphology-based taxonomy and taxon recognition can be problematic in such a large and morphologically highly variable genus; (b) the potential error due to incomplete sampling presents a problem in a genus as variable as Montenegrina; (c) multi-locus analyses should be conducted to arrive at a better basis for species delimitation; and (d) integrative approaches including genetic as well as morphological/anatomical data from a comprehensive geographic sample are necessary.     

Linnaeus' Neptunea (Mollusca: Gastropoda)

Three species of the arcto-boreal, large gastropod Neptunea , described by Linnaeus in 1758 and 1771, occur in large numbers over wide areas of the inshore North Atlantic and adjacent Arctic seas and are conspicuous among Pliocene and Pleistocene molluscs in the Icelandic, North Sea, and western Mediterranean basins. Selections of lectotypes for these species from shells in the collection of the Linnean Society of London, and designations of their type localities, establish the identity of Linnaeus' neptunes and more accurately determine their geographic and geologic distribution. The geographic range of Neptunea (Neptunea) antiqua (L.), the type species, now extends from southern Norway to the northern Biscay coast of France and from the westernmost Baltic Sea to southwestern Ireland; this species also occurs in Pliocene-Holocene marine deposits in West and East Germany, Sweden, the Netherlands, England and France. Its type locality is determined to be the North Sea. N. (Neptunea) despecta (L.) lives in the eastern Canadian Arctic, off southern Greenland, the Barents Sea, and North Atlantic as far south as Massachusetts and Portugal; it also occurs in Pliocene-Holocene strata of eastern Canada, east-central Greenland, Norway (including Svalbard), the Soviet Union, Sweden and England. Its type locality is determined to be the postglacial deposits at Uddevalla in southwestern Sweden. N. (Sulcosipho) contraria (L.) now extends from the southern Biscay coast of France to Cape Spartel, Morocco; this species also occurs in Pleistocene and lower Holocene sequences of the western Mediterranean. Its type locality is determined to be Vigo Bay, Spain. A closely related fossil species, N. (S.) angulata (S. V. Wood), occurs in Pliocene and Pleistocene deposits of the North Sea basin.     

Radiating on oceanic islands: patterns and processes of speciation in the land snail genus Theba (Risso 1826)

Island radiations have played a major role in shaping our current understanding of allopatric, sympatric and parapatric speciation. However, the fact that species divergence correlates with island size emphasizes the importance of geographic isolation (allopatry) in speciation. Based on molecular and morphological data, we investigated the diversification of the land snail genus Theba on the two Canary Islands of Lanzarote and Fuerteventura. Due to the geological history of both islands, this study system provides ideal conditions to investigate the interplay of biogeography, dispersal ability and differentiation in generating species diversity. Our analyses demonstrated extensive cryptic diversification of Theba on these islands, probably driven mainly by non-adaptive allopatric differentiation and secondary gene flow. In a few cases, we observed a complete absence of gene flow among sympatrically distributed forms suggesting an advanced stage of speciation. On the Jandía peninsula genome scans suggested genotype-environment associations and potentially adaptive diversification of two closely related Theba species to different ecological environments. We found support for the idea that genetic differentiation was enhanced by divergent selection in different environments. The diversification of Theba on both islands is therefore best explained by a mixture of non-adaptive and adaptive speciation, promoted by ecological and geomorphological factors.     

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.     

Development of 10 microsatellite loci in the pulmonate snail Biomphalaria kuhniana (Mollusca, Gastropoda)

We characterized 10 variable microsatellite loci in the freshwater snail Biomphalaria kuhniana, as well as conditions for multiplexing and co-loading sets of loci. Two to five alleles were detected per locus over the two studied populations in Venezuela. High inbreeding coefficients suggest high selfing rates. Cross-species amplification provided some variability at eight and three loci in the other species belonging to the Biomphalaria straminea complex (B. straminea and B. intermedia), but was unsuccessful in more divergent species.     

Characterization of 15 microsatellite loci in the pulmonate snail Pseudosuccinea columella (Mollusca, Gastropoda)

We characterized 15 new variable microsatellites in the freshwater snail Pseudosuccinea (Lymnaea) columella, as well as conditions for multiplexing and simultaneously genotyping sets of loci. Two to six alleles were detected per locus over the six populations studied. Gene diversity ranged from 0.000 to 0.498, but essentially no heterozygous individuals were observed. This resulted in extremely high F(IS) estimates, and therefore high selfing rates. The F(ST) estimates ranged from 0.18 to 1 among populations, but was generally high. These markers will constitute efficient tools for investigating the population structure of this invasive species. Cross-species amplification was on the whole unsuccessful.     

Characterization of 17 microsatellite loci in the Japanese land snail genera Mandarina,Ainohelix, and Euhadra (Mollusca,Gastropoda, Pulmonata)

Land snails of the genera Mandarina, Euhadra and Ainohelix are useful for understanding the ecology and evolution of speciation and adaptation, so we have developed 17 microsatellite loci for these species. As in other land snails, most of the loci are highly polymorphic compound repeats, with a great size range between alleles. The loci should be useful in understanding gene‐flow, genetic structure and speciation in these species.