Density-dependent effects in a parasitic nematode,Elaphostrongylus rangiferi,in the snnail intermediate host

Summary Density-dependent effects in Elaphostrongylus rangiferi, a parasitic nematode in the CNS and muscular system of reindeer, were studied in a laboratory population of the snail intermediate host, Arianta arbustorum. The rates in parasite growth, development and mortality were all affected by parasite density. The effects on growth and development were, however, much more marked, than the effect on mortality.All density-dependent rates were intensified by decreasing snail size, and by snail starvation. The snail host showed marked tissue reactions against infection, and the intensity of these reactions increased with increasing parasite density. The mechanism behind the observed density-dependent rates is discussed, and is tentatively concluded to be competition for nutritive substances in the host tissue.The importance of a density-dependent developmental rate in natural populations of this parasite is discussed, and it is hypothesized that this effect may counteract the strong temperature-dependent developmental rate of E. rangiferi In a more general context it is pointed out that density-dependent developmental rates, although common amongst animal populations, has been neglected in models of population dynamics. Developmental rates are usually represented by a constant time lag in such models, but should be treated as a density-dependent variable.     

Snail-type zinc finger proteins prevent neurogenesis in Scutoid and transgenic animals of Drosophila

Scutoid is a classical dominant gain-of-function mutation of Drosophila, causing a loss of bristles and roughening of the compound eye. Previous genetic and molecular analyses have shown that Scutoid is associated with a chromosomal transposition resulting in a fusion of no-oceli and snail genes. How this gene fusion event leads to the defects in neurogenesis was not known until now. Here have found that snail is ectopically expressed in the eye-antennal and wing imaginal discs in Scutoid larvae, and that this expression is reduced in Scutoid revertants. We have also shown that the expressivity of Scutoid is enhanced by zeste mutations. snail and escargot encode evolutionarily conserved zinc-finger proteins involved in the development of mesoderm and limbs. Snail and Escargot proteins share a common target DNA sequence with the basic helix-loop-helix (bHLH) type proneural gene products. When expressed in the developing external sense organ precursors of the thorax and the eye, these proteins cause a loss of mechanosensory bristles in the thorax and perturbed the development of the compound eye. Such phenotypes resemble those associated with Scutoid. Furthermore, the effect of ectopic Escargot on bristle development is antagonized by coexpression of the bHLH gene asense. Thus, our results suggest that the Scutoid phenotype is due to an ectopic snail expression under the control of no-oceli enhancer, antagonizing neurogenesis through its inhibitory interaction with bHLH proteins. Received: 8 February 1999 / Accepted: 24 May 1999     

Impact of invasive apple snails in Hong Kong on wetland macrophytes,nutrients, phytoplankton and filamentous algae

1. Grazing by invasive species can affect many aspects of an aquatic system, but most studies have focused on the direct effects on plants. We conducted mesocosm and laboratory experiments to examine the impact of the invasive apple snail Pomacea canaliculata on macrophytes, filamentous algae, nutrients and phytoplankton. 2. In a freshwater pond, we confined 500 g of Myriophyllum aquaticum or Eichhornia crassipes with 0, 2, 4 or 8 apple snails in 1 m × 1 m × 1 m enclosures for approximately 1 month. Apple snails grazed heavily on both species of macrophytes, with higher overall weight losses at higher snail densities. The damage patterns differed between the two macrophytes. In M. aquaticum, both leaves and stems suffered from substantial herbivory, whereas in E. crassipes, only the roots suffered significant weight reduction. 3. In addition to grazing on macrophytes, apple snails appeared to have controlled the growth of filamentous algae, as these did not develop in the snail treatments. The ability of P. canaliculata to control filamentous algae was supported by a laboratory experiment where the consumption was as high as 0.25 g g⁻¹ snail DW d⁻¹. Because of a lack of native herbivorous snails in the pond, the growth of filamentous algae (mainly Spirogyra sp.) reached 80.3 g m⁻², forming a spongy pond scum in the no‐apple snail control. Together with previous reports that apple snails could eat the juveniles and eggs of other freshwater snails, our results indicated that P. canaliculata could have out‐competed native herbivorous snails from the pond by predation on their juveniles or eggs. Alternatively, P. canaliculata might have out‐competed them by monopolisation of food resources. 4. Nitrogen and phosphorous concentrations remained low throughout both experiments and were not correlated with apple snail density. The treatment effects on chlorophyll a (Chl a) and phytoplankton composition varied in the two experiments. In the M. aquaticum experiment, with increasing snail density, Chl a increased, and the phytoplankton community became dominated by Cryptophyceae. In the E. crassipes experiment, Chl a level was independent of snail density, but with increasing snail density, the phytoplankton community became co‐dominated by Cryptophyceae, Chlorophyceae and Bacillariophyceae. 5. Given the multiple effects of P. canaliculata on wetland biodiversity and function, management strategies should be developed to prevent its further spread. In invaded wetlands, strategies should be developed to eradicate the apple snail and re‐introduce native snails which can control the development of filamentous algae.     

Lead toxicity,locomotion and feeding in the freshwater snail,Lymnaea stagnalis (L.)

The effects of lead (5 or 10 ppm) on the survival of the freshwater snail Lymnaea stagnalis (L.) collected from lead contaminated or uncontaminated environments were evaluated under controlled laboratory conditions. The animals from the contaminated environment had significantly greater survivability than those from the unpolluted environment to subsequent acute (up to 24 days) exposure to lead. Acute (72 h) exposure to lead inhibited several behavioural activities including locomotion, feeding, tentacle extension and emergence from the shell. Lead bioaccumulated in the snail tissues, especially the buccal mass and stomach. The freshwater snail provides a valuable system for studying the bioaccumulation and development of tolerance to environmental lead. Electronic Publication     

Preliminary study on ecology of Bulinus jousseaumei in Schistosoma haematobium endemic rural community of Nigeria

The endemicity of schistosomiasis depends to a large extent on the presence of appropriate freshwater snail species with latent infection, and the quality of the microhabitat of the snails may favour or hinder their development and growth. Monthly in situ determinations of water temperature, pH, total dissolved solid (TDS), conductivity and dissolved oxygen were carried out. Monthly sampling of snails was also conducted. The planorbid snail species morphologically identified were Gyraulus costulatus, Biomphalaria pfeifferi, Bulinus globosus, B. senegalensis, B. jousseaumei, Segmentorbis augustus, Ferrisia sp and Lymnaea natalensis. The most abundant snail species was Gyraulus costulatus (62.2%). Snail density correlated positively with dissolved oxygen (r = 0.349; P = 0.266), while a negative relationship occurred between snail density and conductivity (r = ?0.064; P = 0.843). None of the B. globosus examined shed cercariae, while 12.5% of B. jousseaumei shed cercariae. The occurrence of B. jousseaumei in this study proves its presence in Nigeria and suggests its combined roles with B. globosus in the transmission of urogenital schistosomiasis in the endemic rural communities of Yewa North Local Government Area of Ogun State, Nigeria. Further studies on the geographical distribution of B. jousseaumei are recommended for better understanding of its epidemiological contribution to schistosomiasis in Nigeria.     

Ammonia Excretion in Snails Viviparus bengalensis (LAMARCK) and Pila globosa (SWAINSON) during Active and Dormant Periods

The quantities of ammonia excreted over a period of 24 h by two prosobranch snails Viviparus bengalensis and Pila globosa were measured at 30 day intervals to compare their excretion during the active and dormant phases of their life. The aquatic snail V. bengalensis excreted 0.5-1.0 μmole of ammonia per day and snail throughout the year. The ammonia excretion rate of this snail was 40-50% lower in winter than in the monsoon and summer period. In the amphibious snail Pila globosa excretion of ammonia was limited to only four months of its active life i. e. during July-October. Also the rate of ammonia excretion in P. globosa (0.2-0.3 μmole per day and snail) was very low compared with V. bengalensis.     

The hemolymph of Biomphalaria snail vectors of schistosomiasis supports a diverse microbiome

The microbiome - the microorganism community that is found on or within an organism's body - is increasingly recognized to shape many aspects of its host biology and is a key determinant of health and disease. Microbiomes modulate the capacity of insect disease vectors (mosquitoes, tsetse flies, sandflies) to transmit parasites and disease. We investigate the diversity and abundance of microorganisms within the hemolymph (i.e. blood) of Biomphalaria snails, the intermediate host for Schistosoma mansoni, using Illumina MiSeq sequencing of the bacterial 16S V4 rDNA. We sampled hemolymph from five snails from six different laboratory populations of B. glabrata and one population of B. alexandrina. We observed 279.84 ± 0.79 amplicon sequence variants per snail. There were significant differences in microbiome composition at the level of individual snails, snail populations and species. Snail microbiomes were dominated by Proteobacteria and Bacteroidetes while water microbiomes from snail tank were dominated by Actinobacteria. We investigated the absolute bacterial load using qPCR: hemolymph samples contained 2784 ± 339 bacteria/μl. We speculate that the microbiome may represent a critical, but unexplored intermediary in the snail–schistosome interaction as hemolymph is in very close contact with the parasite at each step of its development.     

Leech predation on juvenile freshwater snails: effects of size,species and substrate

Summary The leechGlossiphonia complanata does not appear to have substantial impact on snail populations, but this may be due to most studies focusing on adult snails rather than juvenile snails. In this study I investigated how predation rates ofG. complanata feeding on newly-hatched and juvenile snails was affected by snail species, snail size, snail density and substrate, in a laboratory experiment. Number of snails eaten increased with increasing density resulting in a type II functional response curve. Predation rates were higher when leeches were feeding onLymnaea emarginata than onPhysa gyrina, whereas there was no significant difference in predation rates when they were feeding onL. emarginata andHelisoma anceps. Sandy substrates and greater snail size resulted in decreased predation rates. Sand reduced movement speed ofG. complanata, which probably reduced encounter rates. Thus, there was a comparatively large effect of leech predation on newly-hatched snails, due to a high probability of encounter and high predation rates, but spatial and temporal refuges probably reduce the importance of leech predation as a structuring force in freshwater snail assemblages.     

Studies on the breeding habits and development of the brood-pouch of a viviparous prosobranch; Melania scabra

Studie son the breeding habits and development of the brood-pouch of a freshwater prosobranch, Melania scabra, have revealed that the snail is ovoviviparous, showing parthenogenetic reproduction. The embryos complete their development in a special incubatory pouch called the brood-pouch or brood chamber, and hatch out in a form completely resembling that of the adult. The snail was found to be a continuous breeder, showing gravid condition throughout the year. General structure, development and functions of the brood-pouch were studied in detail. The brood-pouch was found to contain a slightly alkaline albuminous fluid with a pH 7.2, Qualitative and histochemical studies of the brood-pouch revealed the presence of glycogen, fat and protein as the major constituents. These probably play an important role in the nutrition of embryos.     

Multiple near-identical genotypes of Schistosoma japonicum can occur in snails and have implications for population-genetic analyses

We genotyped (using 16 or 17 microsatellite loci) numerous adult Schistosoma japonicum raised in rabbits exposed to pooled cercariae from small numbers of naturally infected snails from several localities in China. As expected, duplicate multi-locus genotypes (MLGs) were found among these worms. Additionally, many more MLGs, often near-identical, were found than snails used as sources of cercariae. Explanations for these results include (i) genotyping errors, (ii) development within each infected snail of multiple sibling miracidia and (iii) somatic mutation producing genetically varied cercariae from a single miracidium. To control for genotyping errors we re-analysed samples from many individual worms, including repeating the initial PCR. Explanations invoking the development of multiple sibling miracidia within a single snail are not likely to be correct because almost all duplicate MLGs fell within same-sex clusters in a principal coordinates analysis. We would expect both sexes to be represented in a multi-miracidium infection. In addition, we exposed several snails to infection by a single miracidium. One such snail, via an experimentally infected mouse, yielded 48 adult worms. The presence of at least nine near-identical MLGs among these worms was confirmed by re-genotyping. We regard somatic mutation as the most likely explanation for our results. The implications of multiple MLGs for population-genetic studies in S. japonicum are discussed.     

Changes in chemical components in the freshwater apple snail, Pomacea canaliculata (Gastropoda: Ampullariidae), in relation to the development of its cold hardiness

The apple snail, Pomacea canaliculata, is an invasive freshwater snail. It increases its cold hardiness before winter. However, the physiological mechanism of cold hardiness in molluscs is poorly understood, especially in freshwater molluscs. In this study, we examined the changes in low molecular weight compounds, glycogen and lipids, in the body of P. canaliculata in association with the development of cold hardiness. When snails without cold hardiness were experimentally cold-acclimated, the amount of glycerol, glutamine, and carnosine increased, while glycogen and phenylalanine decreased. Overwintering cold-tolerant snails collected from a drained paddy field in November also showed increased glycerol in their bodies with decreasing glycogen concentration, compared to summer snails collected from a submerged field. Water content also decreased during the cold acclimation, although the water loss was minimal. These results indicate that the freshwater snail, P. canaliculata enhances cold hardiness by accumulation of some kinds of low molecular weight compounds in its body as some insects do. However, the actual function of each low molecular compound is still unknown.     

Corallivorous snail removal: evaluation of impact on Acropora palmata

With the continuing decline of Acropora palmata throughout the Caribbean region, impacts of the gastropod corallivore, Coralliophila abbreviata, are becoming more noticeable. A snail removal experiment was performed in remnant A. palmata populations in the Florida Keys National Marine Sanctuary to quantify the area of coral tissue consumed by ambient snail aggregations and to assess the possible effectiveness of snail removal in conserving live coral tissue. Corals where ambient snail aggregations were removed maintained significantly more live tissue area during the 2-month experiment than those where feeding snail aggregations were left in place. The corals with feeding snails left in place lost more than 3 cm² tissue day⁻¹ on average. Thus, removal of C. abbreviata may be an effective measure for conserving depressed A. palmata populations, though secondary effects of such a manipulation remain to be carefully evaluated. Accepted: 26 May 2000     

Save your host,save yourself? Caste‐ratio adjustment in a parasite with division of labor and snail host survival following shell damage

Shell damage and parasitic infections are frequent in gastropods, influencing key snail host life‐history traits such as survival, growth, and reproduction. However, their interactions and potential effects on hosts and parasites have never been tested. Host–parasite interactions are particularly interesting in the context of the recently discovered division of labor in trematodes infecting marine snails. Some species have colonies consisting of two different castes present at varying ratios; reproductive members and nonreproductive soldiers specialized in defending the colony. We assessed snail host survival, growth, and shell regeneration in interaction with infections by two trematode species, Philophthalmus sp. and Maritrema novaezealandense, following damage to the shell in the New Zealand mud snail Zeacumantus subcarinatus. We concomitantly assessed caste‐ratio adjustment between nonreproductive soldiers and reproductive members in colonies of the trematode Philophthalmus sp. in response to interspecific competition and shell damage to its snail host. Shell damage, but not parasitic infection, significantly increased snail mortality, likely due to secondary infections by pathogens. However, trematode infection and shell damage did not negatively affect shell regeneration or growth in Z. subcarinatus; infected snails actually produced more new shell than their uninfected counterparts. Both interspecific competition and shell damage to the snail host induced caste‐ratio adjustment in Philophthalmus sp. colonies. The proportion of nonreproductive soldiers increased in response to interspecific competition and host shell damage, likely to defend the parasite colony and potentially the snail host against increasing threats. These results indicate that secondary infections by pathogens following shell damage to snails both significantly increased snail mortality and induced caste‐ratio adjustments in parasites. This is the first evidence that parasites with a division of labor may be able to produce nonreproductive soldiers according to environmental factors other than interspecific competition with other parasites.     

L'escargot ou les difficultés de l'hermaphrodisme

Summary

The article summarizes the research carried out in Besançon on the stylommatophoran snail Helix aspersa, a pulmonate gastropod mollusc.

The influence that external factors like photoperiod and temperature have on growth and reproduction was studied in this animal. The obtained results made possible a great development of snail breeding.

The potentialities of the hermaphroditism emerge from the observations made on the ovotestis: differentiation and development of the different cell types, presence of fibronectin, richness in neuropeptides, existence of male and female steroids, vitellogenesis processes.

In vivo experiments as well as in vitro cultures demonstrated the involvement of the cerebral complex in the control of reproduction. The dorsal bodies (DB)—(it consists of scattered cells)—stimulate the growth of the oocytes. The DB are under an inhibitory nervous control of the cerebral green cells (CGC); the FMRF-amide might be involved in this process.

A map of the neurons of the cerebral complex has been prepared. It shows the presence and the diversity of neuropeptides. Attention was focused on met-enkephalin and somatostatin. This later peptide seems to be involved in the shell growth. The active molecule, which was isolated from hemolymph and digestive gland is structurally quite similar to vertebrate somatostatin.

The results are compared with those obtained in other molluscs, especially in basommatophora. They reveal that the biological phenomena observed in the snail model are included in the fundamental processes, known in other invertebrates or vertebrates.     

Tissue responses exhibited by Biomphalaria alexandrina snails from different Egyptian localities following Schistosoma mansoni exposure

Snails’ susceptibilities to infection with Schistosoma mansoni were determined through observation of infection rates, total cercarial production and tissue responses of the first generation (F1) of Biomphalaria alexandrina snails, originally collected from different Egyptian governorates (Giza, Fayoum, Kafr El-Sheikh, Ismailia and Damietta) and responses were compared between groups. The emergence of cercariae for a 3-month period and the calculation of survival and infection rates, in control (Schistosome Biological Supply Center; SBSC) and infected snails were evaluated. SBSC and Giza snails showed greater susceptibilities to infection and lower mortality rates. In addition, at 6 and 72 h post-exposure to miracidia all the snail groups showed no difference in the anatomical locations of sporocysts. The larvae were found in the head-foot, the mantle collar and the tentacles of the snails. Sporocysts showed normal development with low tissue reactions in SBSC and Giza snail groups infected with S. mansoni miracidia (SBSC). However, in Fayoum, Kafr El-Sheikh, Ismailia and Damietta snail groups, variable tissue responses were observed in which numerous hemocytes made direct contact with S. mansoni larvae forming capsules. The results suggested that, different responses of B. alexandrina snail’s hemocytes towards S. mansoni are related to the degree of susceptibility of these snails. So this is important in planning the strategy of schistosomiasis control.     

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.     

Competitive interactions between two successful molluscan invaders of freshwaters: an experimental study

The New Zealand mud snail Potamopyrgus antipodarum (Hydrobiidae) and the pulmonate Physella acuta (Physidae) have invaded freshwaters in many parts of the world and become established. They co-exist in many streams, lakes and ponds in New Zealand, often at high densities. In the present study the effects of intraspecific- and interspecific interactions between the two species on growth and reproductive output were examined in laboratory mesocosms. In 30-day experiments, growth of Potamopyrgus antipodarum was lower in high density treatments than controls providing evidence for competition at higher densities of both snail species. No competitive effect was obtained for Physella acuta when controls were compared with high-density treatments, but growth was reduced at high densities of conspecifics. Numbers of juveniles released by Potamopyrgus antipodarum in 40 day trials declined at high snail densities and were lowest at high densities of conspecifics. Egg production by Physella acuta was also reduced at high snail densities. However, when the two species were kept together at equal densities (total snail density twice that of controls), egg production by Physella acuta was significantly higher than in all other treatments, suggesting facilitation by the congenor. Lastly, in a 10-day experiment, Physella acuta grew faster in water conditioned by Potamopyrgus antipodarum than in Physa-conditioned water, whereas Potamopyrgus antipodarum showed no growth response to Physella-conditioned water. Overall, our results indicate that growth and reproductive output of both snail species are influenced more by the density of conspecifics than the presence and density of the other species. The successful co-existence of the two species in New Zealand freshwaters therefore may be a reflection, at least in part, of few competitive interactions between them.     

The distribution of <Emphasis Type="Italic">Biomphalaria</Emphasis> (Gastropoda: Planorbidae) in Lake Victoria with ecological and spatial predictions using Bayesian modelling

Intestinal schistosomiasis is a global disease of enormous public health importance. Assessing the local risk of transmission of the parasite causing the disease requires an appraisal of the distribution of its intermediate snail hosts, Biomphalaria spp.. In East Africa, the Lake Victorian Basin is a major freshwater ecosystem and highly endemic for intestinal schistosomiasis, although detailed distribution data for Biomphalaria have not been collected. We used on-the-ground malacological surveys and conjoint measurement of environmental determinants to develop models for analysis of variables associated with distribution and predictive snail mapping into data-deficient areas. Four expeditions were undertaken to collect snails along the Lake Victoria shoreline, visiting 223 sites overall. Environmental measurements were recorded at the time of inspection and water samples taken for quantification of anion and cation concentrations. The spatial distributions of Biomphalaria choanomphala and Biomphalaria sudanica were modelled in two Bayesian multivariate models: one non-spatial and one spatial with random effects. The results showed that chloride, nitrate, sulphate and the number of sympatric snail species were significant predictors of B. choanomphala whereas habitat, water depth, pH and sulphate were significant predictors for B. sudanica. The range of spatial autocorrelation was large (572.9 km for B. choanomphala and 175.3 km for B. sudanica). Interpolating snail abundance data by kriging revealed two ‘hot-spots’ of high abundance of Biomphalaria. These areas should be targeted in future expeditions to ground-truth the model’s predictions. Our study is the first to use Bayesian methods for determination of biomedically important snail distributions and sets crucial limnological baseline data for assessment of future changes in snail biodiversity in the Lake Victoria Basin.     

Glycosaminoglycans in two mollusks, Aplysia californica and Helix aspersa, and in the leech, Nephelopsis obscura

The presence of glycosaminoglycans was examined in two mollusks (Pulmonates): the terrestrial garden snail, Helix aspersa, and the opishtobranchian sea slug, Aplysia californica and also in the leech (Hirudinea, Erpobdellidae, Nephelopsis obscura). Organs in the garden snail contained predominately chondroitin sulfate and heparan sulfate as a lesser component. The ctenidium of the sea slug contained mainly chondroitin sulfate and a compound which migrated on electrophoresis as heparin but additional data indicated that it could also represent a highly sulfated form of heparan sulfate. The foregut contained only the heparin-like polymer. No standard glycosaminoglycan could be identified in the leech although a polydispersed polysaccharide containing uronic acid, hexosamine and sulfate was shown to be present. A detailed analysis of the heparan sulfate isolated from the garden snail is also given.     

First Record of Podonominae Larvae Living Phoretically on the Shells of the Water Snail Chilina dombeyana (Diptera: Chironomidae / Gastropoda: Lymnaeidae)

Larvae of Podonomus albinervis Edwards, 1931 were found phoretic on Gastropoda Chilina dombeyana (Bruigière, 1789) on the shores of Lake Nahuel Huapi, in Bariloche, Argentina. Larvae were attached with mucus and some had developed a tube made of detritus on the spire of the snail shell. Adults and pupae with associated larval exuviae were found, which helped to identify the species. Approximately 10% of the snail individuals examined on the shores of the lake had chironomid larvae on their shells. Laboratory observations showed that the midge larvae leave the snail shell when the snail leaves the water, thus demonstrating the phoretic relationship. The high density of both snails and Chironomidae in the littoral habitats of the lake may explain this phoretic relationship.