The effects of water availability on the life history of the desert snail,Trochoidea seetzeni

Summary In the Negev desert of Israel, the pulmonate land snailTrochoidea seetzeni is active, grows and reproduces in the month following the torrential winter rains. Thereafter, these snails estivate until the following year's rains. By experimental supplementation of water in the field, we examined the ability of these snails to alter their life histories. Specifically, we measured changes in feeding activity, overall activity levels, movement patterns and reproductive output. We found that snails on artificially-watered plots stopped feeding on higher plants and fed on soil-surface algae growing as a result of increased water availability. Most snails moved onto the watered plots, and reduced the total amount of movement once there, in response to increased food availability. Snails on the watered plots grew more and remained active for a longer period into the summer months than did snails on control plots. However, we found little evidence for the ability of these snails to alter their reproductive outputs in response to increased water (and hence food) availability. There was also no carry-over effect of high water availability in one season enhancing reproduction in the following season. We postulate that the low level of phenotypic plasticity recorded here is a consequence of an adaptive conservative strategy to maintain a high reproductive output under extremely unpredictable desert conditions.     

Distribution and function of an Aplysia cardioexcitatory peptide, NdWFamide, in pulmonate snails

The distribution and function of an Aplysia cardioexcitatory peptide, NdWFamide, were examined in the nervous system of pulmonate snails. We chemically identified the authentic NdWFamide from a land snail (Euhadra congenita) and a freshwater snail (Lymnaea stagnalis). NdWFamide potentiated the heartbeat of those snails. Immunohistochemistry using anti-NdWFamide antibody demonstrated the distribution of NdWFamide-containing neurons and fibers in the central nervous system, as well as peripheral tissues, such as the cardiovascular region and accessory sex organs. These results suggest that NdWFamide is a neuropeptide mediating the neural regulation of the activity of the cardiovascular and reproductive systems of snails.     

Pulmonate phylogeny based on 28S rRNA gene sequences: a framework for discussing habitat transitions and character transformation

Pulmonate snails occupy a wide range of marine, estuarine, freshwater and terrestrial environments. Non-terrestrial forms are supposed to be basal in pulmonate evolution but the group's phylogeny is not well resolved either morphologically or on the basis of available DNA sequence data. The lack of a robust phylogeny makes it difficult to understand character polarization and habitat transformation in pulmonates. We have investigated pulmonate relationships using 27 new sequences of 28S rRNA from pulmonates and outgroups, augmented with data from GenBank. The complete alignments comprised about 3.8kb. Maximum parsimony, maximum likelihood and Bayesian analyses of alignments generated under different assumptions are reported. Complete alignments appear to have a degree of substitution saturation so where there is conflict between hypothesised relationships more weight is given to analyses where regions of random similarity are excluded and which are not affected by this complication. Monophyly of the five main pulmonate groups was robustly supported in almost all analyses. The marine group Amphiboloidea and the freshwater Glacidorbidae are the most basal. The remaining pulmonates (Siphonariidae, Hygrophila and Eupulmonata) form a moderately-supported monophyletic group in all analyses bar one probably affected by saturation of substitutions. Siphonariidae, a predominantly marine and intertidal family, and Eupulmonata (mainly terrestrial with marine, estuarine and freshwater species) form a strongly supported clade that is the sister group to Hygrophila (freshwater). Multiple colonizations of freshwater and terrestrial habitats by pulmonate snails are suggested. No analyses strongly support the possibility of habitat reversions. The colonizations of freshwater by Hygrophila and of land by Stylommatophora were apparently phylogenetically independent although it cannot yet be excluded that there were transient terrestrial phases in the history of the former group or freshwater phases in the latter.     

Ultrastructural investigation of the mechanism of muscle attachment to the gastropod shell

The ultrastructure of the muscle-shell attachment was investigated in the land pulmonate snails Helix aspersa, Anguispira altemata, in the freshwater pulmonate Laevipex sp., and in the freshwater prosobranch Pomacea paludosa. In all cases, a collagenous intercellular matrix and a specialized epithelium (tendon cells) intervene between the columellar muscle and the shell. These tendon cells are characterized by hemidesmosomes at both apical and basal ends, connected by thick bundles of microfilaments. The tendon cells do not insert into the shell directly by microvilli, as formerly thought, but by an extensive network of extracellular organic fibers.     

An evaluation of the interactions between freshwater pulmonate snail hosts of human schistosomes and macrophytes

An account is given of a laboratory investigation designed to evaluate the extent to which the freshwater pulmonate snail Biomphalaria glabrata (Say) can utilize various species of aquatic plants, mainly macrophytes, when presented in the following forms over different time scales: normal plants; dried plant material; homogenized plant material in calcium alginate matrices; water-soluble filtrates of plant homogenates in the medium. The following propositions, derived from the theory of phased coevolution of components of the module consisting of the epiphytic bacteria, algae, snails and macrophytes, are evaluated on the basis of the present results and others including those obtained in this laboratory. That as the snails had become specialized to exploit surface communities of epiphytic algae, decaying plant material and dissolved organic matter (DOM) early in their evolutionary history they would continue to exploit these resources when they later become associated with aquatic macrophytes. That pulmonate snails would tend to be feeding generalists capable of adapting to food of varying chemical composition, given sufficient time, provided it was sufficiently small or flaccid. That although macrophytes and snails show a strong positive relationship, the living macrophyte tissue would be little used by the snails. That the hard outer envelope, inherited from their terrestrial ancestors, would remain as the major defence mechanism of aquatic macrophytes against attack by snails and other aquatic invertebrates. That aquatic macrophytes would invest little in the nutrient deficiency strategy to reduce attack by invertebrates such as snails. That truly aquatic submerged macrophytes would not possess secondary plant compounds (SPC) that would be molluscicidal. Emergent parts of subaquatic or aquatic plants might be expected to be better sources of SPC with molluscicidal factors than submerged aquatic plants. Species of epiphytic or planktonic algae might be better sources of SPC with molluscicidal effects than aquatic macrophytes. That the strategies developed by pulmonate snails for obtaining their energy supplies would not be conducive to rapid speciation. The analysis of the present and other related results supports these propositions. Predictions based on the theory of mutualism involving the snails, macrophytes and other components of the module also receive some support from an analysis of the present results. The additional empirical work that could be undertaken to test this theory is briefly discussed.(ABSTRACT TRUNCATED AT 400 WORDS)     

Native and introduced land snail species as ecological indicators in different land use types in Java

We investigated the effects of different land use types and environmental parameters on the number and abundance of native and introduced land snail species in East Java. 2919 specimens were sampled and assigned to 55 species of which 8 are introduced. Whereas species richness was highest in primary forest, the highest number of introduced species was found in agroforest. The snail assemblages in different habitat types differ much clearer in composition than in total species richness. Plantations and agroforest are dominated by introduced pulmonates with regard to number of individuals, while primary forest is dominated by native prosobranchs. The habitat requirements of the introduced pulmonates differ from those of the native species. In the study area, the abundance of native as well as introduced pulmonate species increased with increasing human impact. However, the abundance of introduced pulmonate species decreased with increasing density of the canopy cover, whereas the abundance of native pulmonate species increased with increasing canopy cover. The abundance of native prosobranch land snails also tends to increase with increasing canopy cover and with the availability of deadwood, but decreased with increasing human impact. Improving the canopy cover and retaining deadwood in plantations and agroforests might help to control the populations of introduced species or even prevent their establishment in these habitats. Land snails are good indicators for the long-term stability of natural habitats, because several species are restricted to undisturbed natural habitats and because of their low dispersal abilities. However, complete inventories of land snail species are costly. Therefore we propose two indices that can be scored with much less effort, namely the percentage of prosobranch individuals and the percentage of individuals of introduced species. Both indices are significantly correlated with the number of native species. Dense plantations and agroforests bordering primary forests may protect the latter from introduced species and help to conserve the native fauna by reducing desiccation and buffering the human impact on the primary forests.     

The feeding habits of Sargochromis Codringtonii in Lake Kariba, Zimbabwe

The food of adult Sargochromis codringtonii in Lake Kariba was dominated by prosobranch snails with pulmonate snails forming an insignificant dietary component. S. codringtonii does not show particular preference for any snail species, feeding on those species that are readily available. Morphometrical analysis of the feeding structures of S. codringtonii showed that the fish is well adapted to handling a molluscivorous diet. The pharyngeal jaws with their flat crowned teeth are well adapted to crushing molluscs. The daily food consumption rate of S. codringtonii was estimated over different seasons. Estimates of food consumption indicate that the fish consumed 13.96% of their dry body weight per day in summer falling to 4.46% in winter. The low consumption of pulmonate snails suggests that S. codringtonii can not be a successful biological control agent against the vectors of schistosomiasis.     

Metabolic rate depression in a marine pulmonate snail: pre-adaptation for a terrestrial existence?

Summary Terrestrial and freshwater pulmonate snails exhibit a marked depression of aerobic metabolism during estivation. This is an adaptation for existence in periodically harsh environments and, though marine gastropods may undergo anaerobic metabolism, they have not been shown to adaptively depress aerobic metabolic rate. We compared the metabolic response to progressive aerial exposure of two intertidal gastropod limpets, a prosobranch and a pulmonate. The prosobranch Patella granularis maintained a constant heart rate until shortly before death. In contrast, the pulmonate Siphonaria oculus underwent facultative depression of heart rate, accompanied by a decline in oxygen consumption. Both heart rate and oxygen consumption returned to normal levels on reimmersion in water. Metabolic rate depression is energy conserving, and may account for the ability of S. oculus to extend higher up the shore than P. granularis, into areas where food availability is low. S. oculus is a primitive, marine pulmonate, periodically subject to harsh conditions, and its capacity for metabolic rate depression may represent a pre-adaptation for life on land.     

Scale-dependence in geographic variation in a freshwater gastropod across the Palearctic

ABSTRACT

The reality of spatial clinal variation in morphological traits of freshwater pulmonate snails (Gastropoda: Pulmonata) has repeatedly been questioned or totally discounted. There is a lack of sound statistical evidence in the articles hitherto published on this subject supporting these claims. Here, by means of different analytical methods (analysis of spatial autocorrelation, linear regression analysis, canonical correlation analysis and others), we demonstrate that shell variation in the dwarf pond snail, Galba truncatula, is patterned in space throughout the northern and central Palearctic, with latitudinally-oriented clines in body size and in some shell proportions. Shell size in G. truncatula decreases with latitude and temperature, representing a special case of converse Bergmann cline. However, the temperature itself is hardly the main driver of shell size variation. It is argued that the shorter growing seasons at high latitudes may represent a better explanation for the observed trend. Shell proportions in the dwarf pond snails vary weakly at the macrogeographic scale, being spatially patterned at lower (mesogeographic) scales around 1200–1500?km. In general, spatial variation in G. truncatula shell size is decoupled from variation in shell shape, demonstrating clear scale-dependence similar to that found in different species of terrestrial (non-aquatic) pulmonate snails.     

An experimental investigation of interactions in snail-macrophyte-epiphyte systems

Summary An experimental investigation under field conditions of enclosures containing freshwater pulmonate snails, the macrophyteCeratophyllum demersum and epiphytes, produced evidence of beneficial interactions.Ceratophyllum growth, measured in terms of stem length, numbers of leaf-nodes and growing tips and leaf survival was significantly enhanced in the presence of snails. This effect was attributed to the increased availability of plant nutrients of snail origin, such as phosphates and ammonia, as well as to the snails' action as “cleaning symbionts” in reducing the density of bacterial and algal epiphyton potentially deleterious to macrophytes. Principal component analysis revealed both seasonal and treatment effects of snail grazing on algal epiphyton. Small adnate algal species (e.g.Cocconeis placentula) survived grazing and benefited from the removal of larger, competitor, species. Snail densities increased in all treatments, despite high (86%) juvenile mortality. It is concluded that freshwater pulmonate snails are strong interactors in lentic habitats, enhancing the growth ofCeratophyllum and producing characterisic epiphyte communities. This benefits not only the snails, but also the plants and epiphytes that are associated with them. Thus the interactions between these component parts of the community can be considered as mutualistic.     

Molecular phylogeny of euthyneura (mollusca: gastropoda)

A new phylogenetic hypothesis for Euthyneura is proposed based on the analysis of primary sequence data (mitochondrial cox1, trnV, rrnL, trnL(cun), trnA, trnP, nad6, and nad5 genes) and the phylogenetic utility of two rare genomic changes (the relative position of the mitochondrial trnP gene, and an insertion/deletion event in a conserved region of the mitochondrial Cox1 protein) is addressed. Both sources of phylogenetic information clearly rejected the monophyly of pulmonates, a group of gastropods well supported so far by morphological evidence. The marine basommatophoran pulmonate Siphonaria was placed within opisthobranchs and shared with them the insertion of a Glycine in the Cox 1 protein. The marine systellommatophoran pulmonate Onchidella was recovered at the base of the opisthobranch + Siphonaria clade. Opisthobranchs, Siphonaria, and Onchidella shared the relative position of the mitochondrial trnP gene between the mitochondrial trnA and nad6 genes. The land snails and slugs (stylommatophoran pulmonates) were recovered as an early split in the phylogeny of advanced gastropods. The monophyly of the Euthyneura (Opisthobranchia + Pulmonata) was rejected by the inclusion of the heterostrophan Pyramidella.     

Molecular systematics of the endemic Leptaxini (Gastropoda: Pulmonata) on the Azores islands

The Azorean representatives of the Leptaxini (Pulmonata) are single island endemics, where a high-spired shell distinguishes the monotypic genus Helixena from two slightly different low-spired forms within Leptaxis (azorica and caldeirarum type). We studied the evolutionary history of putative taxa and the three shell-types using 12 allozyme loci and sequences of nuclear (ITS-1 and ITS-2) and mitochondrial DNA (COI and 16S rRNA). While little variation was found in both ITS genes, allozyme and mtDNA divergence was among the highest reported for pulmonate land snails. Generally, phylogeographic patterns are indicative of allopatric differentiation via the successive colonization of (younger) islands, while a major role for adaptive evolution is not supported. The azorica shell-type is monophyletic and has no common history with other sympatric shell-types on the same islands. The (ambiguous) position of Helixena sanctaemariae makes Leptaxis paraphyletic on the Azores and possibly also the caldeirarum shell-type. Helixena can therefore not be distinguished as a separate genus on the Azores. Following a lineage-based concept, representatives on all (ancient) islands should be considered distinct species.     

Influence of lake trophic status on the occurrence of microcystin-LR in the tissue of pulmonate snails

1. To assess the influence of lake trophic status on the occurrence of microcystin-LR (MCLR) in the tissue of resident pulmonate snails, we sampled seven lakes of varying primary productivity in Alberta, Canada. Parameters associated with productivity were measured every 2 weeks from mid-May through mid-September 1995. Phytoplankton and resident pulmonate snails were collected and analysed for MCLR concentration via high-performance liquid chromatography.
2. For all species of gastropod (i.e. Lymnaea stagnalis , Helisoma trivolvis and Physa gyrina ), the concentration of MCLR in the tissue was correlated ( P  ≤ 0.03) with toxin in the phytoplankton, but not with extracellular aqueous microcystin ( P > 0.28). The concentrations of toxin in the tissues of L. stagnalis and P. gyrina were also correlated with the relative abundance of Microcystis spp. ( P < 0.01).
3. Given that Microcystis spp. abundance and MCLR concentration within phytoplankton are correlated with indicators of productivity, we conclude that trophic status is important in influencing the occurrence and concentration of MCLR in pulmonate snails.     

Polygyrid relations: a phylogenetic analysis of 17 subfamilies of land snails (Mollusca: Gastropoda: Stylommatophora)

The family of polygyrid land snails in North America is significant for its sympatric shell convergences, diversity of mating systems and complex zoogeography; its monophyly and its relation to other families has long been questionable. Cladistic analysis has been performed on one representative each of 17 subfamilies, including all three polygyrid subfamilies and one or more subfamilies each from all ten of the families that have been proposed as the polygyrid sister groups. Eighteen anatomical synapomorphies are used, of which eight are newly discovered, four are differently assessed from previous studies, and six are traditional. The resulting strict consensus tree of alternative maximum-parsimony cladograms is: (Acavidae (Ammonitellidae Corillidae ((Discidae Oreohelicidae) (Helminthoglyptidae Bradybaenidae Polygyridae (Thysanophoridae (Camaenidae Sagdidae)))))).
According to this working hypothesis, the Polygyridae are monophyletic, and their sister group remains unresolved, although the Acavidae, Ammonitellidae, Corillidae, Discidae and Oreohelicidae can be ruled out. Of the five classifications of stylommatophoran families that have been proposed in the past 12 years, the consensus tree is closest to that of Nordsieck. For future morphological work, three regions are recommended as potentially rich in unused phylogenetic information: the fertilization pouch-seminal receptacle complex, the ureter at the pneumostome and the ventral-chain ganglia. Simultaneous dissection, wkh side-by-side comparisons, is recommended over other methods for detecting homologies in land snails. Molecular characters should be exploited, because of the scarcity and the frequent homoplasy of morphological characters.     

Anal uptake of water in terrestrial pulmonate snails

Summary The uptake of water was examined in 35 central European terrestrial pulmonate snail species from 15 families (Table 1) by giving the inactive, withdrawn animals dyed water and observing its bulk flow microscopically.During emergence, 32 of the species studied normally take up water through the pneumostome. A rectal pump (Figs. 1 and 2) rapidly conveys this water through the anus into the gut and stomach.In 7 species, anal water-uptake during emergence was quantified and found to be equivalent to 20–40% of the total body water of the inactive snails before water uptake (Table 2).After emergence, the rectal pump allows re-uptake of urine from the ureter or the lung cativity into the alimentary tract.The anal mechanism permits rapid uptake of large quantities of water into an extrasomal compartment. This helps terrestrial pulmonate snails to exploit short wet spells and presumably helps to stabilize blood concentration during activity under dry conditions.     

Snail predation by larvae of Sepedon scapularis Adams (Diptera: Sciomyzidae), a potential biocontrol agent of snail intermediate hosts of schistosomiasis in South Africa

Malacophagous larvae of the fly Sepedon scapularis Adams were shown experimentally to be effective predators of three species of aquatic pulmonate snails tested as prey: Bulinus africanus (Krauss) an important intermediate host of Schistosoma haematobium (Bilharz), Bulinus tropicus (Krauss) and the invasive species Physa acuta Draparnaud. Survival of S. scapularis larvae from instar to instar was negatively affected by the size of prey snails, since larvae tended to be asphyxiated by the mucous secretions of the snails, or by the larval hydrofuge hairs becoming entangled in snail faeces. In experiments to test the choice of S.scapularis larvae for different species and sizes of snails, B.africanus was significantly disfavoured compared with the other two snail species. Small snails (< 3 mm) were more frequently killed by all three larval instars of S.scapularis. First instars killed few, if any, large snails (> 7 mm), whereas second and third instars preyed effectively on all sizes of snails. Third instars killed significantly more snails than younger instars. For larvae offered only one species of snail as prey, the mean total number of snails killed per larva during its entire development was 49 B.africanus, 45 B.tropicus or 34 P.acuta. It is concluded that S.scapularis is potentially useful as a biological control agent for use against indigenous Bulinus and exotic Physa snails.     

The component community structure of larval trematodes in the pulmonate snail Helisoma anceps

Factors that affected the component community structure of larval trematodes in the pulmonate snail Helisoma anceps in Charlie's Pond, North Carolina, were studied over a 15-mo period using a multiple mark-recapture protocol. Patent infections of 8 species were observed in 1,485 of 4,899 snails examined. Reproductive activity, population size, and survival rate of the snail population were estimated to evaluate the extent of resource availability for the parasites. Antagonistic interactions between trematode species that occurred at the infracommunity level had a neglible effect on the composition and structure of the component community. The patterns observed at this level were related to temporal heterogeneity in the abundance of infective stages (mostly miracidia), differential responses of trematode species to the diverse and constantly changing distribution of snail size and abundance, differential mortality of snails infected with certain trematode species, constant recruitment of 1 trematode species over time, and the existence of predictable disturbances such as the complete mortality of the host population and recruitment of a replacement population during a 6-8 wk period. The last factor operated as a reset mechanism for this snail-trematode system once each year. A model of patch dynamics, with snails as patch resources, best explains the organization and dynamics of this system.     

Cold hardiness abilities vary with the size of the land snail Cornu aspersum

The land snail Cornu aspersum (syn. Helix aspersa) living in Brittany (France) can be considered partially freezing tolerant as it possesses a low ability to supercool and a limited capacity to bear freezing of its body tissues. The absence of a marked cold hardiness strategy permits the emphasis of the role of parameters such as individual size or water mass (W(M)) contained by the organism. Adult snails (shell diameter 30-32 mm) had a supercooling ability, about 1-1.5 degrees C lower than that of immatures (shell diameter 12-20 mm) and survived longer to an exposure to -5 degrees C, with an Lt(50) comprised between 6.0 and 9.8 h against 2.6 to 4.2 h for immature snails. This better ability to bear freezing was explained by the faster dynamic of body ice formation observed in small individuals, which attained ice lethal quantity more rapidly. At the species level, large snails will then tend to be more tolerant to freezing and small ones to be freezing avoidant, a statement also observable at the phylum level.     

Biogeography of the Limacoidea sensu lato (Gastropoda: Stylommatophora): vicariance events and long-distance dispersal

Aim Reconstruction of the historical biogeography of the Limacoidea sensu lato (including the Staffordiidae, Dyakiidae, Gastrodontoidea, Parmacelloidea, Zonitidae, Helicarionoidea and Limacoidea). Evaluation of the relative importance of dispersal and its consequences. Location World‐wide. Methods Weighted ancestral area analysis. Results The ancestral areas of the individual clades have been delimited using weighted ancestral area analysis and a sequence of possible vicariance and dispersal events has been suggested. The results of the ancestral area analysis have tentatively been correlated with Cretaceous and Tertiary palaeogeography. The widely overlapping distribution patterns of several families of the Limacoidea testify to extensive dispersal events. Dispersal capacity of land snails is correlated with body size. The significant negative correlation between body size and distribution area size corroborates the importance of passive dispersal for the evolution of the distribution patterns. Main conclusions The existence of extensive dispersal events of poor active dispersers like land snails diminishes the importance of recent distribution patterns for the reconstruction of palaeogeography. On the other hand, dispersal ensures that biogeographical data reflect the geographical configurations at a given time and renders the use of palaeobiogeographic data for the reconstruction of palaeogeographic configurations of the respective age possible.