Massive upstream migrations by a tropical freshwater neritid snail

While upstream movements of benthic invertebrates have been reported, gregarious upstream migrations of freshwater snails have never been described. We report a possible adaptation to downstream drift by a population of the Neritid (Archeogastropoda) snail,Neritina (Clypeolum) latissima (Broderip, 1833), in the Rio Claro, a small coastal river in Corcovado National Park, Costa Rica. We observed large numbers of these snails in gregarious upstream migration. Snails migrated in narrow lines up to 32 meters long, containing thousands of snails. The migration covered over 1 km of the stream. Larval and juvenile snails are more susceptible to washout by current, and differential mortality of these age classes in the estuarine and ocean environment may provide the selective pressure for the evolution of this behavior.     

Microsatellites in the freshwater snail Bulinus globosus (Gastropoda: Planorbidae) from Zanzibar

Six microsatellite loci were isolated and characterized in Bulinus globosus, a freshwater snail with a wide distribution throughout sub‐Saharan Africa. Bulinus globosus is an intermediate host of Schistosoma haematobium, the causative agent of human urinary schistosomiasis. Microsatellites were tested using 32 snails from four populations collected from Pemba and Unguja islands of Zanzibar. The microsatellite loci displayed relatively low levels of variation, with between two and five alleles per locus. F_ST estimates indicate that gene flow is low, as has previously been suggested for other species of Bulinus.     

Palatability of macrophytes to the invasive freshwater snail Pomacea canaliculata: differential effects of multiple plant traits

1. By selective grazing, invasive grazers can alter macrophyte‐herbivore relationships in shallow freshwater bodies. Evaluating the palatability of macrophytes and understanding the determinants of plant palatability can help predict grazing impact. In no‐choice feeding assays, we tested the palatability of 21 species of freshwater macrophytes to the invasive freshwater apple snail Pomacea canaliculata. 2. Daily feeding rate varied greatly with plant species, ranging from 1.1 to 22% of snail body mass. We assessed six plant properties and examined their correlation with feeding rate. Total nitrogen content was positively related, and C:N ratio and dry matter content (DMC) negatively related, to snail feeding rate. There was no significant correlation between snail feeding rate and plant phenolic content, but the feeding rate on Myriophyllum aquaticum (the plant with the highest phenolic content) was very low. 3. We repeated the feeding assays for 15 species that were not palatable as fresh leaves with reconstituted plant tissues formed by mixing ground up dried leaves with agar. The feeding rate still differed greatly among macrophyte species. Phragmites australis and Vallisneria natans (two species with the highest DMC) were eaten much more as reconstituted plant than as fresh leaves, indicating that structure (i.e. DMC) may be important in their defence against snail herbivory. For two plants (M. aquaticum and Alternanthera philoxeroides) that had moderate amounts of nitrogen/phosphorus but were consumed very little as fresh and reconstituted tissues, we incorporated their extracts into a palatable agar‐based food. The extracts from both species greatly reduced snail feeding rate, indicating the presence of chemical defences in these two species. 4. These results indicated that feeding was affected by several plant traits. The snail favoured plants with a high nitrogen content and avoided plants with a high DMC. Only a few plants possessed chemical feeding deterrents that were effective against this snail. Given the invasive spread of P. canaliculata in Asia, ecologists and managers should consider plant palatability when selecting plants for use in wetland restoration and when predicting the impact of further invasion by this species.     

Body size-dependent gender role in a simultaneous hermaphrodite freshwater snail, Physa acuta

We examined whether gender role in the simultaneous hermaphroditefreshwater snail, Physa acuta, is determined by relative bodysize in a manner predicted by the size-advantage model. We observedthe body-size combinations of pairs in the laboratory by usingfield-collected populations. Smaller individuals tended to playthe "male" role (sperm donor), and larger snails the "female"(sperm recipient). Next, we analyzed the mating behaviors involvedin gender-role decision in snail pairs of three different body-sizecombinations, using "large" and "small" snails. Smaller snailswere more likely to approach the partner as a male in different-sizecombination (large/small), whereas frequent initial approachesas a male and rejection behavior as a female were observed inthe large/large combination. Third, we examined the body sizepreference when a snail can freely choose the partner from twoother individuals of different body sizes (large/large/smallor large/small/small). Small individuals had a significant tendencyto act as the male and positively selected large snails as thefemale partner in both triple combinations. However, the largeindividual acted as both the male and the female with nearlyequal frequency. In the size-differing pairings, copulationsoccurred after fewer male approaches and fewer rejections thanin pairings involving two large snails, suggesting that bodysize difference is one of the behavioral solutions in genderconflict. Clear gender-role switching associated with body sizewas not seen. Smaller snails thus have a tendency to play themale role more frequently but adopt both gender roles when theirbody size is sufficiently large.     

Difference in habitat selection by different morphs of the land snail Arianta arbustorum

The behaviour of morphs of Arianta arbustorum has been investigated using a new technique involving a paint which fades in daylight. The technique was used to study the response of different morphs to exposure to sunlight at a homogeneous woodland habitat and a heterogeneous open habitat. The yellow morph showed a significantly higher rate of fading than the brown morph. A mark, release and recapture programme was used to analyse the association between frequencies of different morphs recaptured and not recaptured. In both habitats, there was a general tendency for the frequency of recapture to increase with increasing age in both morphs. Habitat selection resulting from different behaviour of different morphs of A. arbustorum is discussed. Morph frequencies can be related to habitat, the brown morphs of A. arbustorum are significantly less frequent in the open habitat. The behavioural responses of Cepaea hortensis , present with A. arbustorum in two other populations, to the exposure of sunshine were also studied. Climatic selection by solar energy associated with different morphs, species and habitat are also discussed.     

Litter shredding in a desert oasis by the snail Melanopsis praemorsa

The freshwater snail Melanopsis praemorsa is veryabundant inoases of the Judean Desert. To determine its role as littershredder, experiments were carried out in Nahal Arugot. Boxeswereplaced in the stream (29 °C), each containing 1 g ofdrywillow leaves together with one snail; or six snails.With one snail per box, dry leaf weight after two daysdeclined to72% of the initial dry weight, after twelve days to 58% andaftereighteen days to only 40%. During this period thesnails in the experiment increased by 12% in shell-height, by20%in snail weight (including the shell), by 33% in live freshbiomass. Each snail consumed about 10 mg litter per day.With six snails per box, dry leaf weight after two daysdeclined to62% of the initial dry weight and after twenty days to only15%.In natural populations where density was 44 snails per quadrat(of25×25 cm), 1 g of dry leaves was reduced to 25%theinitial weight within three days. Overall plant cover along a600×10 m stretch of the stream was found to be45%, ofwhich 65% was willow. During one year about 1 kg litter wasshedper m² willow; by extrapolation, this would be about1800 kglitter throughout the stretch. Much of this vast amount oflitteris consumed by the snails, who thereby reduce summer dammingof theoasis. Our study suggests that M. praemorsa, a generalistfeeder, may be a major litter shredder in certain desert oasisecosystems.     

Hybridization and invasiveness in the freshwater snail Melanoides tuberculata: hybrid vigour is more important than increase in genetic variance

Many invasive taxa are hybrids, but how hybridization boosts the invasive process remains poorly known. We address this question in the clonal freshwater snail Melanoides tuberculata from Martinique, using three parental and two hybrid lines. We combine an extensive field survey (1990-2003) and a quantitative genetic experiment to show that hybrid lines have outcompeted their parents in natural habitats, and that this increased invasiveness co-occurred with pronounced shifts in life-history traits, such as growth, fecundity and juvenile size. Given the little time between hybrid creation and sampling, and the moderate standing genetic variance for life-history traits in hybrids, we show that some of the observed trait changes between parents and hybrids were unlikely to arise only by continuous selection. We therefore suggest that a large part of hybrid advantage stems from immediate heterosis upon hybridization.     

Inheritance of isozyme phenotype at three loci in the freshwater snail,Goniobasis proxima: Mother-offspring analysis and an artificial introduction

A good deal of interest has recently focused on the genetics of Goniobasis populations, with isozyme frequencies often a primary tool. I sampled 72 Goniobasis proxima from a population naturally polymorphic at three enzyme loci, Gpi, Opdh, and Est, reared each in isolation, and electrophoretically analyzed 14 sibships produced. Since females apparently mate but once in their lifetimes, it was possible to infer paternal genotype and demonstrate that inheritance is Mendelian at these three loci. The three loci appear to assort independently. A pair of G. proxima populations fixed for alternative alleles at the three loci has been introduced into a small spring. Triply heterozygous F₁ hybrids have been recovered, although at a low frequency.A grant from College of Charleston R & D funds is gratefully acknowledged.     

The influence of self‐fertilization and grouping on fitness attributes in the freshwater snail Physa acuta: population and individual inbreeding depression

The genetic structure, selfing rate and inbreeding depression of the hermaphroditic freshwater snail Physa acuta were jointly analysed in a population near Montpellier, France. Allozymic markers revealed moderate gene diversity (0.138), and no heterozygote deficiency. The mean outcrossing rate, estimated by using progeny arrays, was 0.9, with substantial variation among families. This also suggests that the number of fathers among outcrossed offspring of a given mother is low. Inbreeding depression was estimated over more than one generation using 83 first‐laboratory‐generation (G₁) families. The main parameters measured were parental (G₁) fecundity, offspring (G₂) survival and fecundity. Size and growth were also monitored. Parental fecundity was analysed under several conditions (isolation, pair and quadruplet outcrossing). The self‐fertilization depression, including parental fecundity, offspring survival and fecundity, was about 0.9 at the population level. The genetic data obtained in the same population indicate a value of about 0.3 using Ritland’s (1990) technique, suggesting that the depression over the whole life‐cyle might be even higher than 0.9. Grouping affected neither fecundity nor self‐fertilization depression. Substantial variation in depression for survival was detected among individuals, from no survival in some selfed families to better survival than that of outbred families in others. The overall result (outbred population structure, high outcrossing rate and high self‐fertilization depression) is consistent with what is expected in large outcrossing populations in which inbreeding depression is maintained by mutation‐selection balance.     

Leaf mechanical properties modulate feeding movements and ingestive success of the pond snail, Lymnaea stagnalis

We examined the mechanical properties of Butterhead and Iceberg lettuce leaves, and the rate at which they were eaten by the pond snail Lymnaea stagnalis. The outer part of Butterhead leaves were less robust than either the inner Butterhead or outer Iceberg leaves (Young’s modulus 2.8, 5.2, 7.7 MPa respectively; ultimate tensile stress 0.18, 0.34 0.51 MPa) which were also thicker. Snails ingested inner Butterhead and Iceberg strips more slowly (36 and 32%) than outer Butterhead. This was not due to differences in latency to first bite or biting rate. Rather, the drop was due to a decrease in the proportion of successful bites (inner Butterhead 84%; Iceberg 86%), to a shorter length ingested per bite (inner Butterhead 55%; Iceberg 45%) and to increased handling time (inner Butterhead 30%). We conclude that sensory input from the mechanically more robust lettuce slows the buccal central pattern generator.     

First record of the freshwater snail Pseudosuccinea columella (Gastropoda: Lymnaeidae) in southern Pampas (Argentina) and assessment of future spread

ABSTRACT

The freshwater snail Pseudosuccinea columella was recorded for the first time in Argentina more than 60 years ago. Since then its distribution in the wild has been restricted to the northeastern provinces. Here we record the presence of P. columella in southern Pampas for the first time, extending its distribution more than 500?km southwards. The climatic suitability of this and other areas of South America for its establishment and spread was analysed using habitat modelling software. Hitherto its spread within and between watercourses in southern Pampas has been very limited, probably through a combination of low climatic suitability, recent introduction and low connectivity of the drainage basins. The suitability of other areas where it has been recently recorded indicates a moderate risk of further spread in central and northwestern Argentina and in coastal areas of the Pacific rim of South America (southern Perú and northern Chile). The recent spread of P. columella in the wild in Argentina may be the result of an increase in the trade in aquarium plants or of the evolution of a new lineage with different ecological capabilities.     

Clonal diversity driven by parasitism in a freshwater snail

One explanation for the widespread abundance of sexual reproduction is the advantage that genetically diverse sexual lineages have under strong pressure from virulent coevolving parasites. Such parasites are believed to track common asexual host genotypes, resulting in negative frequency‐dependent selection that counterbalances the population growth‐rate advantage of asexuals in comparison with sexuals. In the face of genetically diverse asexual lineages, this advantage of sexual reproduction might be eroded, and instead sexual populations would be replaced by diverse assemblages of clonal lineages. We investigated whether parasite‐mediated selection promotes clonal diversity in 22 natural populations of the freshwater snail Melanoides tuberculata. We found that infection prevalence explains the observed variation in the clonal diversity of M. tuberculata populations, whereas no such relationship was found between infection prevalence and male frequency. Clonal diversity and male frequency were independent of snail population density. Incorporating ecological factors such as presence/absence of fish, habitat geography and habitat type did not improve the predictive power of regression models. Approximately 11% of the clonal snail genotypes were shared among 2–4 populations, creating a web of 17 interconnected populations. Taken together, our study suggests that parasite‐mediated selection coupled with host dispersal ecology promotes clonal diversity. This, in return, may erode the advantage of sexual reproduction in M. tuberculata populations.     

Crayfish predation on the common pond snail (Lymnaea stagnalis): the effect of habitat complexity and snail size on foraging efficiency

Optimal foraging theory was used to explain selective foraging by the introduced signal crayfish (Pacifastacus leniusculus) on the thin-shelled common pond snail (Lymnaea stagnalis). Crayfish predation efficiency was studied in relation to habitat complexity and snail size. In a pool experiment (area 1.3 m²) single adult crayfish were allowed to feed on four size classes of snails for one week. A pair-wise preference trial (aquarium experiment) tested if adult crayfish selectively predated on particular size classes of snail and if prey value (expressed as snail dry mass per handling time) could explain the size range of snails chosen. Crayfish preferred the smallest size classes of snails in both pool and aquaria experiments. In the pool experiment crayfish had a strong effect on snail survival. Habitat complexity did not affect overall snail survival, but resulted in reduced predation pressure on the smallest size classes of snails. Handling time and shell-thickness increased exponentially with increasing snail size, and the two smallest size classes had the highest prey values. The results suggest that crayfish can structure the abundance and size distribution of thin-shelled snails, through size-selective predation and reduction of macrophytes. The mechanisms behind the choice of snails may be based on prey value and reduced exposure time to predators and conspecifics. Crayfish effects on snail size distribution may be less pronounced in complex habitats such as macrophyte beds. This revised version was published online in July 2006 with corrections to the Cover Date.     

Temperature adaptation in the freshwater snail, helisoma trivolvis (say), in an artificially heated reservoir in the southeastern United States

Snails from a heated zone of Par Pond of the U.S. Savannah River Plant were compared with conspecifies from an unheated area of the same reservoir. The heated area averaged 5°C warmer and was thermally more variabic than the control area. This situation has existed for about twenty years. Variation in metabolic response to temperature was observed according to season and heated vs unheated area. A laboratory rearing experiment indicated that part of the observed variation was environmetally induced.

Life cycle and growth rates were similar in the two collection areas. Three generations were produced annually in both areas and spawning was synchronous except that the summer spawning period was shorter in the heated area. Live weights expressed relative to shell size were also similar, and decreased in both areas during summer.

It appears from these results that metabolic compensation for temperature can serve as an indicator of general thermal adaptability, but only if the degree of environmental plasticity is known for the species.     

A comparative analysis of metacommunity types in the freshwater realm

Most metacommunity studies have taken a direct mechanistic approach, aiming to model the effects of local and regional processes on local communities within a metacommunity. An alternative approach is to focus on emergent patterns at the metacommunity level through applying the elements of metacommunity structure (EMS; Oikos, 97, 2002, 237) analysis. The EMS approach has very rarely been applied in the context of a comparative analysis of metacommunity types of main microbial, plant, and animal groups. Furthermore, to our knowledge, no study has associated metacommunity types with their potential ecological correlates in the freshwater realm. We assembled data for 45 freshwater metacommunities, incorporating biologically highly disparate organismal groups (i.e., bacteria, algae, macrophytes, invertebrates, and fish). We first examined ecological correlates (e.g., matrix properties, beta diversity, and average characteristics of a metacommunity, including body size, trophic group, ecosystem type, life form, and dispersal mode) of the three elements of metacommunity structure (i.e., coherence, turnover, and boundary clumping). Second, based on those three elements, we determined which metacommunity types prevailed in freshwater systems and which ecological correlates best discriminated among the observed metacommunity types. We found that the three elements of metacommunity structure were not strongly related to the ecological correlates, except that turnover was positively related to beta diversity. We observed six metacommunity types. The most common were Clementsian and quasi‐nested metacommunity types, whereas Random, quasi‐Clementsian, Gleasonian, and quasi‐Gleasonian types were less common. These six metacommunity types were best discriminated by beta diversity and the first axis of metacommunity ecological traits, ranging from metacommunities of producer organisms occurring in streams to those of large predatory organisms occurring in lakes. Our results showed that focusing on the emergent properties of multiple metacommunities provides information additional to that obtained in studies examining variation in local community structure within a metacommunity.     

Influence of water temperature on the activity of the freshwater snail Pomacea canaliculata (Caenogastropoda: Ampullariidae) at its southernmost limit (Southern Pampas,Argentina)

1.

The effect of temperature on the activity of the freshwater snail Pomacea canaliculata was investigated through field surveys and laboratory trials.     

Microsatellite variation in the freshwater snail Physa acuta

We present data for seven new markers isolated from a microsatellite‐enriched DNA library from the freshwater snail Physa acuta. Six of them appear to be conserved in Physa cubensis. These loci might be useful for analysing population structure and reconstructing the story of invasion of freshwaters on a worldwide scale by P. acuta.     

Delayed spermatophore removal in the land snail Euhadra peliomphala

Although many studies have demonstrated that variation in male genital morphology can influence male fertilization success, it remains unclear in many cases which mechanism(s) of sexual selection and sexual conflict produce the morphological divergence of genitalia. Distinguishing between such mechanisms requires, at the very least, information on the process by which genital morphology influences fertilization success. The length of the spermatophore‐producing organ (SPO) of the stylommatophoran land snails is a case in point. The length of the spermatophore may have an important influence on the transfer of the spermatophore to the recipient's genital tract. To test this hypothesis, we investigated in Euhadra peliomphala (Pfeiffer, 1850) the relationship between the length of the SPO in sperm donors and the position of their spermatophore in sperm recipients at 72, 84, and 96 h after mating. The results showed that longer spermatophores required longer times to reach a gametolytic organ and to begin to be digested. These findings suggest that this delay in spermatophore digestion explains the mechanism for the high fertilization success shown by snails with a long SPO. © 2013 The Linnean Society of London