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Despite their ubiquity and considerable biomass, the roles played by parasites in aquatic food webs are still not well understood, especially those of their free-living infectious stages. For instance, cercariae, the motile larvae of parasitic flukes (trematodes) may be a key source of nutrients and energy for consumers. As cercariae clonally reproduce within the digestive-gonadal gland complex of gastropod intermediate hosts that acquire nutritionally important polyunsaturated fatty acids (PUFA) mainly from their diets (e.g., by grazing on primary producers), cercariae could transfer snail-derived PUFA if consumed. Through fatty acid (FA) analysis, we explored whether a change in the diet of parasitized hosts altered the FA profiles of both snail-only and trematode-containing snail tissue, thereby affecting their nutritional values. Freshwater snails (Stagnicola elodes) infected with Plagiorchis sp. were fed three different diets (cyanobacteria, green algae, and diatoms) that differed in nutritional quality with respect to FA profiles. While diet influenced the overall FA composition of both snail-only tissue and snail tissue containing trematodes, levels of certain PUFA (mainly omega-3) were largely unaffected. Trematode-containing snail tissue also generally contained more PUFA relative to snail-only tissue. Notably, both tissue types had far higher levels of PUFA than found in their diets. Our results suggest that freshwater snail hosts, and possibly their associated trematode parasites, could be trophic upgraders of key PUFA despite anthropogenically induced changes in algal communities that may lead to overall diminished PUFA contents. As such, cercariae-mediated trophic transfers of PUFA may play important roles in aquatic food webs.

Understanding the extent to which traits that are used to delimit and diagnose species are phenotypically plastic is important for recognizing species boundaries. Shell characters have long been used for describing species of gastropods, even though these features may be influenced by environmental conditions. To determine the degree of phenotypic plasticity of two North American lymnaeid species, Stagnicola elodes (Say, 1821) and Stagnicola emarginata (Say, 1821), that occur in different habitats and differ in shell morphology, we reared individuals in captivity under similar conditions and compared shell shapes and sizes of wild-caught and captive-reared populations. We also exposed individuals of S. elodes to effluent from a potential predator (crayfish) to gauge the possible impact of the presence of predators on shell morphology. Although the two species remained morphologically distinct, shell shapes of captive-reared individuals of both species differ significantly from those of wild-caught individuals and show similar magnitudes of change among species. Directions of change, however, differed significantly among species. Although shell shapes of individuals of S. elodes that were exposed to crayfish cues were not significantly different from control snails, shell sizes of exposed snails were smaller than unexposed snails. These results suggest that exposure to predators affects growth rates of S. elodes. Nonetheless, given significant associations between shell shape and size that were observed in the captive-rearing and predator-exposure experiments, shell shape changes allometrically during development. These results suggest that morphological differences of other North American Stagnicola species reflect ecophenotypic variation, but more work is necessary to further evaluate this hypothesis.

Trematode-induced castration of snails is widespread and can lead to other life history changes of snails such as changes in trajectories of size and growth or survival. The changes produced likely depend on whether the parasite or host controls allocation of host resources remaining after partial or complete cessation of host current reproduction by castrating trematodes. Documenting host life history changes, like changes in host size in response to castration, is a first step in assessing whether these changes are beneficial to the parasite (increasing transmission success) or to the host (outliving the infection) or to neither. Herein, we test for differences in size and survival among individuals of two snail species in relation to infection by Echinostoma spp. trematodes. Active shedding of Echinostoma spp. was associated with castration of all Stagnicola elodes snails from a site in Eastern Ontario. Snails actively shedding cercariae were not different in size from non-shedding, egg-laying snails but had a higher mortality than egg-laying snails. Active shedding of Echinostoma spp. cercariae was also associated with castration of nearly all Helisoma trivolvis monitored, from a site in Southwestern Ontario. Actively shedding, non-laying H. trivolvis hosts were smaller on average than non-shedding egg-laying hosts, but both non-laying and egg-laying snails survived equally well. We discuss these results in light of what is known about effects of castration on snail hosts in terms of growth and survival for these and other trematode species and speculate on whether changes in size or survival benefits parasite or host.

In this study, we provide the first molecular evidence for a possible connection between freshwater mollusc faunas across the Bering Strait via the Beringian Land Bridge using data inferred from gastropods of the family Lymnaeidae. The gastropods collected from geothermal springs in the Tumrok Mountains, West Kamchatka, Russia, share the nuclear internal transcribed spacer 2 (ITS2) and the cytochrome oxidase subunit I gene (COI) haplotypes, thus being as sister to those recorded for lymnaeid snails in the Stagnicola elodes group from Canada and the USA. Two lymnaeid species, Lymnaea (Orientogalba) tumrokensis Kruglov and Starobogatov, 1985 and Lymnaea (Polyrhytis) kurenkovi Kruglov and Starobogatov, 1989, were described from the Tumrok geothermal locality, but actually they are morphological variations of a single taxon of subspecies rank re-classified here as Ladislavella catascopium tumrokensis. This subspecies is the first discovered representative in the genus, which formed a dwarf race in a geothermal habitat. Our findings highlight the possible exchange between freshwater faunas in Beringia during the Pleistocene and an important role of geothermal ecosystems as possible cryptic refugia for freshwater hydrobionts.

Environmental stressors have the potential to greatly impact the transmission of parasites with complex, multi-host life cycles such as those of trematodes. The commonly used herbicide atrazine has been shown to affect the susceptibility of second intermediate hosts (such as larval amphibians) to trematode infection, as well as the longevity and infectivity of the free-swimming cercariae, but not eggs or the free-swimming miracidia that infect the gastropod first intermediate hosts. However, we do not know if this pesticide influences the survival of infected snails or whether it affects cercariae production within, or emergence from, these hosts. In addition, previous studies of host-parasite dynamics have only examined the parent atrazine compound, not any of the long-lasting metabolites commonly present in water bodies. Here, we report that a concentration of 0.33 µg/L of an atrazine metabolite, desethyl atrazine, increased the mortality of freshwater gastropods ( Stagnicola elodes ) infected with a gymnocephalus type of cercaria but not that of uninfected snails or those harboring a mature or dormant infection of Echinoparyphium sp. In contrast, 2 wk of exposure to desethyl atrazine did not affect the emergence of gymnocephalus cercariae from snails, although a trend for a decrease in the emergence of Echinoparyphium sp. cercariae was observed. We suggest that simultaneous trematode infection and exposure to contaminants may represent a significant combined stress to gastropods, but this is likely parasite species-specific as well as dependent on whether cercariae are being actively produced.

Despite the importance of litter process- ing in aquatic ecosystems, we have an incomplete knowledge of roles that specific benthic invertebrate taxa play in leaf decomposition. We studied the effect of gastropods on leaf litter decay rates in two outdoor pond mesocosm experiments: one contrasted the individual effects of six gastropod taxa on decompo- sition of senescent red maple (Acer rubrum) leaves and another evaluated the effect of Stagnicola elodes on decomposition of senescent leaves from five common deciduous forest tree species. Decay rate of red maple leaves was significantly higher with gastropods present. Rates increased by 14-39% in the presence of the relatively large-bodied pulmonates S. elodes and Helisoma trivolvis, while the small- bodied pulmonate Physa gyrina and the prosobranchs Goniobasis livescens and Bellamya japonica had smaller, non-significant effects. S. elodes significantly increased the decay rates of black cherry (Prunus serotina), sugar maple (Acer saccharum), red maple, and red oak (Quercus rubra) leaves by 72, 41, 34, and 25%, respectively, relative to no gastropod controls. Our results suggest that some gastropod species may be important players in the processing of allochtho- nous detritus, particularly in smaller, forest-enclosed lentic systems where leaf litter inputs constitute a large portion of the energy budget.

Experiments on the toxicity of cadmium (Cd(2+)) to the embryonic development of Stagnicola elodes (Mollusca, Gastropoda), obligatory first intermediate host of numerous trematodes of pathogenic importance, were carried out as part of a study on the effects of metal pollution on host-parasite relationships. Freshly laid snail eggs were exposed to Cd concentrations of 0, 0.02, 0.2, and 2.0 mg Cd(2+)/L, and survival and embryogenesis were examined for 30 days. Mean survival time (+/- SD) of the control group was 23.1 (+/- 5.3) days compared with 10.1 (+/- 3.2) at 0.02 mg Cd(2+)/L, 3.9 (+/- 0.7) at 0.2 mg Cd(2+)/L, and 1.1 (+/- 0.08) at 2.0 mg Cd(2+)/L. Mortality patterns of all test groups differed significantly from each other, demonstrating that the percentage of surviving individuals at any given time was inversely related to Cd concentration. Concentration-dependent effects of Cd exposure on snail embryogenesis were noted. While embryos of the control group developed properly and started hatching on day 16, eggs exposed to 0.02 mg Cd(2+)/L exhibited a prolonged gastrula period and failed to hatch. Eggs in the 0.2 mg Cd(2+)/L group were blocked in the gastrula stage on day 5, whereas individuals exposed to 2.0 mg Cd(2+)/L died in the morula stage on the second day. Data showed that Cd severely affects S. elodes embryogenesis. By implication, Cd contamination at concentrationsor=0.02 mg Cd(2+)/L will thus decrease transmission success of various trematodes by decreasing intermediate host snail abundance.

Experimental and field studies suggest that freshwater snail species have negative effects on each other's population growth rates. Because snails share similar diets, these interactions have been interpreted as the result of exploitative competition, but they could also result from intraguild predation. Here we present three experiments aimed at testing the hypothesis that interspecific interactions among three species of freshwater gastropod (Helisoma trivolvis, Physa acuta and Stagnicola elodes) are mediated by intraguild egg predation. Foraging trials, conducted in a laboratory, showed that some snails readily prey on eggs, but the extent of egg predation depended on both the identity of the snail predator and the identity of the egg mass. Of the three species considered, Stagnicola had the largest effect on egg mortality and Physa had no effect on egg mortality. Foraging trials also showed that the eggs of Physa were the most vulnerable to predators and that the eggs of Stagnicola were largely invulnerable. A study conducted in large outdoor mesocosms assessed the occurrence of egg mortality in an environment of more extensive spatial scale and complexity. The results largely mirrored those of the laboratory study, with Stagnicola being the most voracious predator and the eggs of Physa being most vulnerable to predation. The reproductive success of Physa and Stagnicola raised in sympatry and allopatry was assessed in a mesocosm study conducted over three months. Recruitment of both species was depressed in sympatry, but patterns of growth in the survivors suggest contrasting mechanisms of suppression: Physa suppressed Stagnicola via exploitative competition, but Stagnicola suppressed Physa via egg predation. These experiments support the hypothesis that freshwater snail assemblages are structured by strong interspecific interactions and that a rich interplay of egg predation and interspecific competition underlie interactions among the members of this guild.

Studies in lakes show that fish and crayfish predators play an important role in determining the abundance of freshwater snails. In contrast, there are few studies of snails and their predators in shallow ponds and marshes. Ponds often lack fish and crayfish but have abundant insect populations. Here we present the results of field surveys, laboratory foraging trials, and an outdoor mesocosm experiment, testing the hypothesis that insects are important predators of pulmonate snails. In laboratory foraging trials, conducted with ten species of insects, most insect taxa consumed snails, and larval dragonflies were especially effective predators. The field surveys showed that dragonflies constitute the majority of the insect biomass in fishless ponds. More focused foraging trials evaluated the ability of the dragonflies Anax junius and Pantala hymenaea to prey upon different sizes and species of pulmonate snails (Helisoma trivolvis, Physa acuta, and Stagnicola elodes). Anax junius consumed all three species up to the maximum size tested. Pantala hymenaea consumed snails with a shell height of 3 mm and smaller, but did not kill larger snails. P. acuta were more vulnerable to predators than were H. trivolvis or S. elodes. In the mesocosm experiment, conducted with predator treatments of A. junius, P. hymenaea, and the hemipteran Belostoma flumineum, insect predators had a pronounced negative effect on snail biomass and density. A. junius and B. flumineum reduced biomass and density to a similar degree, and both reduced biomass more than did P. hymenaea. Predators did not have a strong effect on species composition. A model suggested that A. junius and P. hymenaea have the largest effects on snail biomass in the field. Given that both pulmonate snails and dragonfly nymphs are widespread and abundant in marshes and ponds, snail assemblages in these water bodies are likely regulated in large part by odonate predation.

Atrazine has been shown to affect freshwater snails from the subcellular to community level. However, most studies have used different snail species, methods, endpoints, and atrazine exposure concentrations, resulting in some conflicting results and limiting our understanding. The goal of this study was to address these concerns by (1) investigating the acute and chronic effects of atrazine on four species of freshwater snails (Biomphalaria glabrata, Helisoma trivolvis, Physa acuta, and Stagnicola elodes) using the same methods, endpoints, and concentrations, and (2) summarizing the current literature pertaining to the effects of atrazine on freshwater snails. We conducted a 48 h acute toxicity test with an atrazine concentration higher than what typically occurs in aquatic environments (1000 µg/L). Additionally, we exposed snails to environmentally relevant atrazine concentrations (0, 0.3, 3, and 30 µg/L) for 28 days and assessed snail survival, growth, and reproduction. We also summarized all known literature pertaining to atrazine effects on freshwater snails. The literature summary suggests snails are often affected by environmentally relevant atrazine concentrations at the subcellular and cellular levels. These effects are typically not transitive to effects on survival, growth, or reproduction at the same concentrations. Our acute exposures corroborate the general trend of no direct effect on snail populations as atrazine did not directly affect the survival of any of the four snail species. Similarly, environmentally relevant concentrations did not significantly affect the survival, growth, or reproduction of any snail species. These results indicate that, in the absence of other possible stressors, the direct effects of environmentally relevant atrazine concentrations may not be realized at the snail population level.

A panel of 4 digenetic trematode species (Echinostoma paraensei, E. trivolvis, Schistosoma mansoni, and Schistosomatium douthitti) and 5 snail species (Biomphalaria glabrata, Helisoma trivolvis, Lymnaea stagnalis, Stagnicola elodes, and Helix aspersa) was examined to determine if known patterns of host specificity could be explained by the tendency of digenean larvae to be bound by snail hemocytes, or by the ability of larvae to influence the spreading behavior of hemocytes. In short-term (1 hr) in vitro adherence assays, there was no overall pattern to suggest that sporocysts were more likely to be bound by hemocytes from incompatible than compatible snails. Compared with the other parasites, sporocysts of E. paraensei were less likely to be bound by hemocytes from any of the snail species tested. All rediae examined, including those of another species Echinoparyphium sp., were also remarkably refractory to binding by hemocytes from any of the snails. Of all the larvae examined, only sporocysts and young daughter rediae of E. paraensei caused hemocytes to round up in their presence. This was true for hemocytes from the compatible species B. glabrata and the incompatible lymnaeid species S. elodes and L. stagnalis. The patterns of host specificity shown by this particular panel of parasites and snails were not predicted by either the extent of hemocyte adherence to digenean larvae or by the ability of larvae to affect hemocyte spreading behavior. The results of this study suggest that a role for hemocytes, although likely, may require different assays, possibly of a more prolonged nature, for its detection. Also, different parasite species (notably E. paraensei) and intramolluscan stages have distinctive interactions with host hemocytes, suggesting that the determinants of specificity vary with the host-parasite combination, and with the parasite life cycle stage.

The enzyme β-hydroxybutyrate catalyzes the interconversion of the ketone bodies, acetoacetate and β-hydroxybutyrate. Although the activity of β-hydroxybutyrate dehydrogenase is relatively high in freshwater and terrestrial molluscs, it is undetectable in all marine molluscs examined. We tested the effects of conditions of tissue preparation and ionic strength of assay media on the measurement of β-hydroxybutyrate dehydrogenase and conclude that the apparent absence of this enzyme in marine molluscs is not an artifact of conditions of measurement. Measurements of other enzymes involved in ketone body and lipid metabolism in the hepatopancreas of the marine snail Littorina littoria , the freshwater snail Stagnicola elodes and the terrestrial snail Cepaea nemoralis suggest that the differential expression of the enzyme may be related to differences in the metabolic organization of marine molluscs and their freshwater and terrestrial counterparts. Mapping of the presence/ absence of β-hydroxybutyrate dehydrogenase onto a bivalve and gastropod phylogeny indicates four independent occurrences of the enzyme.

The host response to and the viability and infectivity of metacercariae of Cyclocoelum mutabile were examined in three species of pulmonate snails, Stagnicola elodes, Gyraulus parvus, and Promenetus exacuous. Host response was generally similar among snail species at 8 weeks post infection. Host responses increased with age of infection in S. elodes. There was no difference in the excystment success of metacercariae from each species at 8 weeks post infection. The mean excystment time in vitro was greater for metacercariae from S. elodes than for those from either G. parvus or P. exacuous at 8 weeks post infection. There were no differences in either excystment success or mean excystment time of metacercariae from 5-, 8-, and 30-week-old infections from S. elodes. The mean infection success of metacercariae from S. elodes was lower than that of metacercariae from G. parvus and P. exacuous in juvenile coots. Adult coots were less susceptible to infection than juveniles. There was no difference in the infection levels established in naive and previously infected adult coots.

The herbicide atrazine is a photosynthetic inhibitor used around the world in agricultural applications. Contamination of surface waters adjacent to treated areas can directly reduce growth of nontarget aquatic autotrophs, but the severity of impacts is highly dependent on species sensitivity and exposure concentration. Secondary effects resulting from macrophyte or phytoplankton decline may include an expansion of the more tolerant periphyton community. Recently, this shift in the autotrophic community has been proposed as a mechanism for increased rates of parasite infections in amphibians via augmented populations of aquatic snails which act as intermediate hosts to larval trematodes. To further clarify this relationship, an outdoor microcosm study was conducted to examine the effects of atrazine on primary production and snail populations over a range of environmentally relevant concentrations. In July 2009, 15 experimental ponds were treated to achieve initial concentrations of 0, 1, 10, 30, and 100 µg/L atrazine. Over a period of 73 d, measures were taken of macrophyte, phytoplankton, and periphyton biomass, growth, and fecundity of caged snails (Physella spp. and Stagnicola elodes) and free-living snails (Physella spp.). Except for declines in macrophyte biomass at the highest treatment level, no consistent relationships were found between atrazine concentration and any measured parameter. Comparison of these results with previous findings highlights the variability of responses to atrazine exposure between similarly constructed freshwater communities, even at concentrations up to 20 times higher than sustained environmental levels.

During the period from January to December (2007), 1,095 freshwater molluscs of four species were captured (Lymnaea stagnalis, Stagnicola elodes, Physella cubensis and Physa acuta) in Lake Chicnahuapan, State of Mexico, Mexico. Two hundred seventy-two (24.84% prevalence) of these molluscs were parasitised by 11 trematode species (from which two were not identified at the species level) having six cercariae species and five metacercariae species represented in five families. The cercariae Telorchis corti (Plagiorchiidae) and the metacercariae Cotylurus cornutus (Strigeidae) were the species with the highest prevalence among the examined snails. The highest percentage of infection was observed in L. stagnalis (27.45% of prevalence, n = 572) and P. cubensis (23.96%, n = 455). Twenty-one of the examined snails had multiple infections with up to three trematode species.

The life cycle of Sciomyza varia (Coquillett) is described. The species is behaviorally and morphologically specialized for breeding in stranded or aestivating Stagnicola elodes (Say) (Gastropoda: Lymnaeidae), an aquatic snail. Eggs are laid on the host's shell in a consistent pattern. The solitary, parasitoid larvae feed selectively on host tissues, and the host remains alive during most of larval development. The distinctive puparia are highly modified to fit inside the host's shell and occlude the aperture. Data on the habitat, behavior, feeding habits, and phenology of adults and immature stages are presented, and the Nearctic distribution is mapped. Certain features of the egg, three larval instars, and puparium are illustrated and described. Evolution of specialized malacophagy is discussed, and the immature stages are compared with those of other Sciomyzidae.

SUMMARY. 1. The impact of crayfish predation on the abundance of macroinvertebrates was examined under semi-natural conditions. Female (Experiment 1) or male (Experiment 2) crayfish (Orconectes virilis) were held for 5 weeks in twelve small pools (4.67 m2 surface area) at biomasses of 0. 5, 10 or 18 g m−2 (live weight). The pools were stocked with known densities of macroinvertebrates. 2. Crayfish significantly affected the abundance of macroinvertebrates in the pools. Differences in the effects of crayfish on macroinvertebrates were related to crayfish sex, the presence of age-0 crayfish, and the species of macroinvertebrate. 3. The abundance of snails (Stagnicola elodes and Physa gyrina) was greatly reduced, in comparison with controls, by biomass of female crayfish ≥10 g m−2 and by biomasses of male crayfish ≥5 g m−2. The total density of non-molluscan invertebrates was inversely correlated with the biomass of female crayfish but the total biomass of non-molluscan invertebrates did not differ between treatments. This is consistent with our observation that small invertebrates (

Allatostatin-like immunoreactivity was demonstrated in the central nervous system (CNS) neurons of the freshwater pulmonate snails Bulinus globosus (Planorbidae) and Stagnicola elodes (Lymnaeidae). Immunopositive neurons were localized using a monoclonal antibody highly specific to allatostatin I (APSGAQRLYGFGL-amide) from the cockroach Diploptera punctata. All CNS ganglia in both snail species contained immunopositive neurons. The pedal ganglia showed large numbers (80-100) of neurons in Bulinus and Stagnicola. Large numbers of immunopositive cells were also found in the cerebral (60) and buccal ganglia (20) of Bulinus. Other ganglia contained fewer immunopositive cells, but these cells were most concentrated in the cerebral (Stagnicola) and left parietal (Bulinus) ganglia and the visceral ganglia of both species. The high concentration of immunopositive cells in the pedal ganglia and axons demonstrable in the pedal nerves suggests that one possible function for a molluscan allatostatin-like peptide would be to modulate muscular function. Extract of Stagnicola CNS effected 50% inhibition of juvenile hormone synthesis by corpora allata of D. punctata at between 15 and 30 CNS equivalents, providing further evidence that the molluscan immunoreactive material is a peptide, or peptides, with sequence similarity to the active part of the D. punctata allatostatins.

The herbicide Rodeo® provides waterfowl managers with an effective chemical tool for creating open water habitats in wetlands if its use does not adversely affect native invertebrate communities. The survival of caged Chironomus spp. (midge), Hyalella azteca (amphipod), Stagnicola elodes (pond snail), and Nephelopsis obscura (leech) was assessed in prairie pothole wetlands treated by air with a tank mixture of Rodeo®, the surfactant X-77 Spreader®, and the drift retardant Chem-Trol® at a rate recommended for controlling cattails. Laboratory studies were then conducted to determine the acute toxicities of Rodeo®, X-77 Spreader®, and Chem-Trol®, individually and in simulated tank mixtures, to the same invertebrates and to Daphnia magna in reconstituted water representative of these wetlands. There was no difference in the survival of caged invertebrates between treated and reference wetlands after 21 days. Based on nominal concentrations of the formulations, X-77 Spreader® (LC50s=2.0–14.1 mg/L) was about 83–136 times more toxic than Rodeo® (LC50s=218–1216 mg/L) to aquatic invertebrates. Chem-Trol® killed ≤10% of the animals at 10,000 mg/L and ≤50% of the animals at 28,000 mg/L. Daphnia magna were more sensitive than the other species to X-77 Spreader®, Rodeo®, and the simulated Rodeo® tank mixture (RTM). The joint toxic action of the RTM was additive for amphipods and midges, greater than additive for leeches, and was less than additive for daphnids. X-77 Spreader® was the major toxic component in the RTM. Binary mixtures of X-77 Spreader®, Rodeo®, and Chem-Trol® at tank mixture and equitoxic ratios also showed additive toxicity to amphipods. The use of Rodeo® (applied as a tank mixture with X-77 Spreader® and Chem-Trol®) as a management tool in wetlands does not pose an acute hazard to native aquatic invertebrates because the concentrations of Rodeo®, X-77 Spreader®, and Chem-Trol® found to be acutely toxic to these invertebrates were much higher than their expected or measured concentrations in water from wetlands treated with the RTM.

SUMMARYWe investigated changes in the behaviour of Stagnicola elodes associated with the emergence of Plagiorchis elegans cercariae. Within 15 min of the reduction in light intensity, which triggered the onset of cercarial emergence, infected snails moved to the top of the water column and remained there for 2–3 h. Seventy-nine percent of all cercariae that emerged from the snail did so during this period. Uninfected snails showed no such behavioural changes following the change in light intensity. Cercariae were released in a dense cloud around the snail at the water surface and dispersed passively. Within 3–4 h more than 80% of all cercariae had settled in the bottom 5 cm of the water column. The infectivity of cercariae increased from less than 20% upon emergence from the snail to greater than 75% 4–6 h post-emergence, and then declined steadily to below 5% by 24 h post-emergence. Cercarial longevity was greater than 30 h and exceeded the period of infectivity. This may be related to steadily falling endogenous glycogen levels. A delay in attaining maximum infectivity may represent an adaptive mechanism allowing time for cercarial dissemination, thus reducing superinfection, and subsequent parasite-associated mortality, of second intermediate hosts.

Digenetic trematodes usually show a high degree of specificity for their molluscan intermediate hosts. A panel of 4 digenean species (Echinostoma paraensei, E. trivolvis, Schistosoma mansoni, and Schistosomatium douthitti) and 5 snail species (Biomphalaria glabrata, Helisoma trivolvis, Lymnaea stagnalis, Stagnicola elodes, and Helix aspersa representing 3 gastropod families) was used to assess the relative contributions of miracidial behavior, host plasma osmolality, and host plasma factors in dictating specificity. Additional experiments were undertaken with a fifth digenean, Echinoparyphium sp. Expected patterns of compatibility were first confirmed; each parasite species produced patent infections in its known snail host, but not in the other snail species. One exception was S. douthitti, which unexpectedly did not infect L. stagnalis. As judged by direct observation and by noting their disappearance after exposure to snails, miracidia were generally less likely to attach to or penetrate incompatible than compatible hosts. However, over half of the miracidia of each parasite species attached to or attempted penetration of both compatible and incompatible hosts, suggesting that under the experimental conditions used, miracidial host location and attachment behaviors were not of overriding importance in dictating observed patterns of specificity. For each digenean species, the percentage of larvae that became immobile, rounded, showed tegumental damage, or died over a 6-hr interval in plasma of the various snails was assessed. In no case was plasma from a compatible host harmful to sporocysts or rediae. In contrast, in 8 of 16 (50%) incompatible combinations, snail plasma had a significant negative effect on sporocyst condition. In 4 of 12 (33%) incompatible combinations, plasma had a significant negative effect on rediae. In 9 of 10 combinations tested, lymnaeid plasma was toxic for the parasites of planorbid snails and in 2 of 4 combinations, planorbid plasma was toxic for the parasites of lymnaeid snails. Toxicity was not attributable to differences in plasma osmolality between snail species. The ability of plasma from incompatible snails to affect viability of both sporocysts and rediae was surprisingly strong, suggesting that humoral factors play a greater role in dictating patterns of digenean-snail specificity than previously appreciated.

Eggs of Plagiorchis elegans were readily ingested by Stagnicola elodes of all ages, but were more infective to immature than mature snails. Infection enhanced the growth of the host in a dose-dependent manner. The number of cercariae released by immature snails increased with the age of the snail host; mature snails yielded fewer cercariae. Heavily infected snails tended to die prematurely, thereby reducing their total production of cercariae to levels below those of more lightly infected individuals. Even light infections castrated the snail host, Snails that acquired the infection as juveniles never produced eggs. Actively reproducing snails ceased egg laying within days of infection and never recovered. All parasite effects on the growth and reproduction of the snail host first manifested themselves during the early stages of infection, long before the development of daughter sporocysts and cercariae, and are therefore attributable to the effects of mother sporocysts. The study provides insight into how this natural enemy of mosquito larvae may be established in natural snail populations by means of strategically timed introductions of parasite eggs, with a goal of maximizing cercarial production for the biological control of sympatric mosquito larvae.

number of eggs available over the course of incubation (egg days) (219.00 ? 2.24) remained low. These values increased progressively as incubation temperatures rose and reached levels of 3.59 ? 0.30 and 1,467.50 + 4.56, respectively, at 20 C. Although incubation at 30 C further increased the mean number of eggs per day (5.45 ? 0.56), the mean number of egg days declined sharply to 735.25 ? 4.71, suggesting that elevated temperatures enhanced embryonation but lowered the survival of embryonated eggs. This was also reflected in the infectivity of eggs over time. Embryonated eggs incubated at 10 and 20 C remained infective significantly longer than eggs incubated at higher or lower temperatures. Incubation in constant light yielded significantly lower mean numbers of embryonated eggs per day (1.86 + 0.09) and mean number of egg days (96.25 ? 0.99) than incubation in constant darkness (2.23 + 0.17 and 701.50 ? 2.86 eggs, respectively) but did not affect the infectivity of embryonated eggs. The data suggest that at moderate temperatures and shielded from light, fully embryonated eggs of P. elegans may survive in the aquatic environment for a period of months. Such eggs may serve as a source of infection for populations of Stagnicola elodes and ensure the sustained production of the highly entomopathogenic cercariae required for mosquito control. Entomopathogenic digeneans have been proposed as agents in the biological control of mosquitoes (Rao et al., 1985; Rau, 1992). Preliminary field trials have shown that the introduction of Plagiorchis-infected snails into mosquito habitats reduces adult mosquito emergence and longevity (Rau et al., 1991). Where mosquito preimagos share their aquatic environment with the molluscan first intermediate host, Stagnicola elodes (Say, 1821), this digenean infection may be introduced by deploying the eggs of the parasite. Infective eggs measuring 32-41 X 17-24 ,xm (StyczynskaJurewicz, 1962) are ingested by the snail host, penetrate the intestinal wall, and form mother sporocysts in its tissues. Daughter sporocysts are produced asexually and these, in turn, give rise to the entomopathogenic cercariae. The xiphidiocercariae leave the snail host and, after a brief free-swimming existence, penetrate the tissues of aquatic insect larvae, among them mosquitoes, in which they form metacercariae. They are transmitted to the avian or small mammal definitive host by ingestion of infected insects. Metacercariae excyst in the intestinal tract of the definitive host and grow into adults. Eggs are passed with the host's feces and develop to infectivity in the external environment (Styczynska-Jurewicz, 1962; Genov and

Infection with the digenean parasite Plagiorchis elegans dramatically reduced the reproductive output of Biomphalaria glabrata exposed to the parasite as juveniles or adults. The total number of eggs produced by infected snails was reduced to approximately 7 and 13% of control values, respectively. Parasitic castration was attributed to the presence of mother sporocysts that readily established in the tissues of this incompatible host. Infection did not result in the production of cercariae but significantly shortened the life span of juvenile and adult B. glabrata by approximately 23 and 10%, respectively. Plagiorchis elegans also castrated its compatible host, Stagnicola elodes.

Infectivity of Cyclocoelum mutabile miracidia to 9 species and up to 4 size classes of pulmonate snails at 14, 16, and 20 C was studied under laboratory conditions. Of the 9 species examined, 6 (Stagnicola elodes, Lymnaea stagnalis, Gyraulus parvus, Gyraulus circumstriatus, Promenetus exacuous, and Armiger crista) were highly susceptible (infection success > or = 25%), 2 (Physa jennessi and Helisoma trivolvis) had low susceptibility (infection success 0), and 1 (Physa gyrina) was not susceptible to infection. Within highly susceptible species, snail size was negatively related to susceptibility and temperature had variable effects. Infection success was not affected by temperature or snail size in species with low susceptibility. Production of cercariae was negatively correlated with susceptibility among snails of different sizes and species, but was not influenced by snail size for a given species. Among species, metacercariae production was typically higher in lymnaeids than in either planorbids or physids. Results of experiments where miracidia were provided with a choice of 2 different snails suggest that they do not discriminate between species with high and low susceptibility.

The transmission of Cyclocoelum mutabile to snails was examined under natural conditions by sampling the snail communities of 4 natural ponds that had been exposed experimentally to infection by laboratory-infected coots (Fulica americana). Five of 6 snail species in the ponds, Physa jennessi, Promenetus exacuous, Armiger crista, Gyraulus parvus, and Stagnicola elodes, became infected. No natural infections were found in the few Helisoma trivolvis examined. The second most abundant species Promenetus exacuous was infected most often, whereas Physa jennessi, the most abundant species present, was rarely infected. The temporal pattern of infections in the snail community suggests the transmission window of this parasite in southern Manitoba is limited by both the 14 C hatching threshold of the fluke eggs and the seasonality of ovigerous infections in the coot host. No naturally overwintering infections were found in snails from these ponds, which were examined the following spring. None of the 1,120 laboratory-infected snails placed in cages and held overwinter in the ponds survived, whereas 14 of the 1,120 uninfected control snails kept in the same cages survived. These results are consistent with the hypothesis that C. mutabile must be reestablished in northern waterfowl breeding areas each spring.

Sixteen species of aquatic snails of four families were tested by quantitative technique under standardized conditions for their suitability as intermediate hosts for Angiostrongylus cantonensis. These species were the planorbid snails Biomphalaria glabrata, Biomphalaria alexandrina, Planorbis planorbis, Planorbis intermixtus, Bulinus truncatus, Bulinus contortus, Bulinus africanus, Bulinus tropicus and Helisoma sp.; the lymnaeid snails Lymnaea natalensis, Lymnaea tomentosa, Lymnaea stagnalis, and Stagnicola elodes; the physid snail Physa acuta (an Egyptian and a German strain) and the ampullariid snails Marisa cornuarietis and Lanistes carinatus. All these snail species proved to be susceptible to infection with A. cantonensis, and first stage larvae reached the infective third stage in all of them. However, the rate and intensity of infection varied with different species. B. glabrata was the most susceptible snail species with a 100% infection rate and an average percentage recovery of third stage larvae of 26.1. This was followed by S. elodes and B. africanus, with a 100% infection rate and an average percentage recovery of third stage larvae of 15.6 and 14.6 respectively. The rest of snail species proved to be less susceptible. For comparative evaluation of the suitability of the various snail species as intermediate hosts of A. cantonensis a "Capacity Index" was determined. This index should provide a useful method for the evaluation of the suitability of various snails as intermediate hosts of nematode parasites under standardized conditions in the laboratory.

SummaryThe inheritance of alleles at ten enzyme loci of Stagnicola elodes (Lymnaeidae) was analysed by laboratory crosses. Alleles for alanine aminopeptidase (2 alleles), leucine aminopeptidase (2 alleles), aspartate aminotransferase (2 alleles), β-hydroxybutyrate dehydrogenase-1 (4 alleles), 6-phosphogluconate dehydrogenase (2 alleles), esterase-2 (2 alleles), esterase-3 (2 alleles), esterase-7 (2 alleles), esterase-13 (2 alleles) and esterase-18 (2 alleles) segregate in the F2 generation according to Mendelian expectations. Contingency X2 tests for linkage done on F2 progeny from crosses between strains homozygous for alternate alleles at two loci showed that β-hydroxybutyrate dehydrogenase-1, esterase-13 and esterase-18 are presumably linked.

Pulmonate and prosobranch gastropods collected in Minnesota and Washington were examined for the presence of coccidian parasites. Typical coccidian oocysts were recovered from the feces of 89 of 543 (16%) pulmonate snails representing the families Lymnaeidae, Physidae, Planorbidae, and Succineidae. No coccidial infections were apparent in 104 specimens representing the prosobranch families Viviparidae and Pleuroceridae. Oocysts identified as Pfeifferinella ellipsoides were recovered from 21 of 101 (21%) Stagnicola elodes, 2 of 94 (2%) Physa gyrina, and 4 of 67 (6%) Aplexa hypnorum although incidence of infection differed greatly (2–40%) dependent upon the locality sample. This coccidian has not been described previously from North America and is reported here from members of the families Lymnaeidae and Physidae for the first time. Twenty percent of Oxyloma retusa from one habitat were shedding oocysts slightly smaller than, but otherwise identical with, those of Alveocystis gugleri. A third as yet unidentified oocyst was recovered from up to 54% of Helisoma trivolvis. As a group, the coccidia of pulmonate molluscs have been poorly studied. The findings of this survey, however, suggest that these parasites may be more common and widely distributed in both the Basommatophora and Stylommatophora than previously believed.

Young pond snails of three species (Lymnaea stagnalis, Stagnicola elodes, and Helisoma trivolvis) were allowed free access to waterfowl feathers floating on the surface of shallow containers. Snails adhering to the feathers were subjected to simulated flight conditions by placing them in an air stream. The size of these snails, the length of time they adhered to the feathers while in the air stream, and their survivability for various lengths of time under these conditions were recorded. Those snails that adhered to the feathers were mainly small individuals (

The vertical distribution of juvenile pond snails (Lymnaea stagnalis and Stagnicola elodes), hatched from egg masses containing known numbers of viable embryos, was recorded daily in artificial containers under outdoor conditions for up to 25 days after hatching. Both species showed an initial vertical movement to the surface of the water with a subsequent return to the bottom sediments after 10–15 days. This pattern was consistent despite differences in the depth of the water column (13 versus 18.5 cm). At 25 days after hatching more than 85% of the population was in the lower half of the container at any one time, almost all being associated with the bottom sediments. The presence of an adult snail of either species in the same container significantly reduced the survival of young snails (from between 85 and 94% to between 1 and 18%). Consumption of the young snails during their first 10 days out of the egg mass, when they were moving to the surface, accounted for the observed differences in survival. Young snails appeared to be ingested incidently by the foraging adults, most often when the former were concentrated at the air–water–glass interface.

The resource allocation strategies of the freshwater snails Stagnicola elodes and Physella gyrina were examined using artificial diets diluted to various degrees with cellulose. Even when nutrients were diluted by 50 or 75%, both species showed remarkable maintenance of their rates of assimilation (94 to 294% of controls) and production (82 to 140% of controls). This was mainly achieved by varying rates of feeding (148 to 397% of controls) and respiration (76 to 839% of controls). S. elodes was twice as large as P. gyrina and had slower rates. The hypothesis that this endows S. elodes with greater compensatory scope (i.e., a greater range of response in feeding or respiration rates) was strongly supported. This observation has broad implications for theories of evolution and niche breadth. Particular attention was devoted to trade-offs between somatic and reproductive sup- port. With 75% dietary dilution, the two species showed exactly inverse strategies. S. elodes supported growth more than reproduction (90 and 59% of controls, respectively), whereas P. gyrina supported reproduction over growth (116 and 38% of controls, respectively). Their strong compensatory ability suggests that these snails normally operate at submaximal rates that are homeostatically maintained. Shifts in resource allocation may be engaged as "anticipatory" safety measures, even before absolute shortages arise.

The suitability of Fossaria (Bakerilymnaea) cubensis and Pseudosuccinea columella from Louisiana as intermediate hosts for Fasciola hepatica was compared to P. columella and Stagnicola elodes from Ann Arbor, MI, S. attenuata from Hidalgo, Mexico, Lymnaea gedrosiana from Iran and L. natalensis from Senegal. P. columella from LA was shown to be a more suitable host (51.3% became infected) than F. (B) cubensis (15.2% and 26.4% of two populations became infected). The infection rate for P. columella from MI, was 50% and for L. gedrosiana was 32.5%, whereas L. natalensis, S. elodes and S. attenuata were refractory. F. (B) cubensis and P. columella have some degree of suitability as intermediate hosts for F. hepatica under laboratory conditions, but field observations of their habitat in southern LA and characteristic management of cattle indicate that the former snail is more important as an intermediate host in this enzootic area. Tissue sections of suitable snails had few histopathological effects but physical damage caused by rediae was pronounced, mainly in the digestive gland and in the mantle.