Predation and the distribution and abundance of a pulmonate pond snail

Summary The abundances of a freshwater pulmonate snail, Lymnaea elodes were studied in a temporary pond and a permanent, more productive pond in northeastern Indiana, USA. When snails from both populations were reared in each of the ponds in containers excluding predators, snails grew to be 1.3 to 2 times as large in the more productive pond, and laid 9 times as many eggs. However, field sampling data showed adults to be more abundant in the temporary pond. The only obvious difference between the two ponds was the presence of the molluscivorous central mudminnow (Umbra limi) in the permanent pond. These fish fed upon L. elodes when eggs and juvenile snails were abundant. In an experiment in the temporary pond, addition of mudminnows lowered egg and juvenile snail survival in pens where snail abundances had been increased. We suggest that vertebrate predators like the mudminnow can be significant sources of mortality for thin shelled species like L. elodes, possibly excluding them from habitats like lakes and rivers.Address for offprint requests     

Hydroperiod,predators and the distribution of physid snails across the freshwater habitat gradient

1. Studies of species distributions across environmental gradients further our understanding of mechanisms regulating species diversity at the landscape scale. For some freshwater taxa the habitat gradient from small, shallow and temporary ponds to large, deep and permanent lakes has been shown to be an important environmental axis. Freshwater snails are key players in freshwater ecosystems, but there are no comprehensive studies of their distributions across the entire freshwater habitat gradient. Here we test the hypothesis that snail species in the family Physidae are distributed in a non‐random manner across the habitat gradient. We sampled the snails, their predators and the abiotic environment of 61 ponds and lakes, spanning a wide range in depth and hydroperiod. 2. Temporary habitats had the lowest biomass of predators. Shallow permanent ponds had the highest biomass of invertebrate predators but an intermediate fish biomass. Deep ponds and lakes had the highest fish biomass and intermediate invertebrate biomass. Five species of physids occurred in the regional species pool and 60 of the 61 ponds and lakes surveyed contained physid snails. Each pond and lake contained an average of just 1.2 physid species, illustrating limited membership in local communities and substantial among‐site heterogeneity in species composition. 3. Physids showed strong sorting along the habitat gradient, with Physa vernalis found in the shortest hydroperiod ponds and Aplexa elongata, P. gyrina, P. acuta and P. ancillaria found in habitats of successively greater permanence. When organised into a site‐by‐species incidence matrix with sites ordered according to their hydroperiods, we found the pattern of incidence to be highly coherent, showing that much of the heterogeneity in species composition from one pond to another is explained by hydroperiod. We also found that the number of species replacements along this gradient was higher than random, showing that replacement is more important than nesting in describing species composition in ponds of different hydroperiod. 4. Discriminant analysis showed that pond depth, invertebrate biomass and fish biomass were the best predictors of species composition. Analysis of these niche dimensions showed that P. vernalis and A. elongata were most successful in shallow, temporary ponds with few predators. P. gyrina and P. acuta were typically found in ponds of intermediate depth and high predator abundance. P. ancillaria was found in the deepest lakes, which had abundant fish predators but few invertebrate predators. Of the five species considered, P. ancillaria, P. vernalis and A. elongata were relatively specialised with regard to key habitat characteristics, P. gyrina was moderately generalised and P. acuta was remarkably generalised, since it alone occurred across the entire freshwater habitat gradient. The exceptional habitat breadth of P. acuta stands in contrast to distributional studies of other freshwater taxa and deserves further attention.     

COSTS AND BENEFITS OF A PREDATOR-INDUCED POLYPHENISM IN THE GRAY TREEFROG HYLA CHRYSOSCELIS

The phenotypes of gray treefrog (Hyla chrysoscelis) tadpoles vary depending on whether predators are present in the pond. Tadpoles reared in ponds with predatory dragonfly larvae are relatively inactive compared with tadpoles in predator-free ponds, and have relatively large, brightly colored tailfins with dark spots along the margins. Models for the evolution of plasticity predict that induced phenotypes such as this should confer high fitness relative to the typical phenotype when in the presence of predators, but should be costly when the predator is absent. Our study tested for the predicted fitness trade-off in H. chrysoscelis by first rearing tadpoles in mesocosms under conditions that induce the alternate phenotypes, and then comparing the performance of both phenotypes in both environments. We generated the two phenotypes by rearing tadpoles in 600-liter outdoor artificial ponds that contained either two caged dragonflies (Anax junius) or an empty cage. Tadpoles from the two environments showed significantly different behavior, tail shape, and tail color within two weeks of exposure. We compared the growth and survival of both phenotypes over four weeks in ponds where there was no actual risk of predation. Under these conditions, both phenotypes grew at the same rate, but the predator-induced phenotype had significantly lower survival than the typical phenotype, indicating that induced tadpoles suffered greater mortality from causes other than odonate predation. We tested the susceptibility of both phenotypes to predation by exposing them to dragonflies in 24-h predation trials. The predator-induced phenotype showed a significant survival advantage in these trials. These results confirm that the predator-induced phenotype in H. chrysoscelis larvae is associated with fitness costs and benefits that explain why the defensive phenotype is induced rather than constitutive.     

Competition and intraguild egg predation among freshwater snails: re-examining the mechanism of interspecific interactions

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.     

Prey responses to fine‐scale variation in predation risk from combined predators

While it is well documented that organisms can express phenotypic plasticity in response to single gradients of environmental variation, our understanding of how organisms integrate information along multiple environmental gradients is limited in many systems. Using the freshwater snail Helisoma trivolvis and two common predators (water bugs Belostoma flumineum and crayfish Orconectes rusticus), we explored how prey integrate information along multiple predation risk gradients (i.e. caged predators fed increasing amounts of prey biomass) that induce opposing phenotypes. When exposed to single predators fed increasing amounts of prey biomass, we detected threshold responses; intermediate amounts of consumed biomass induced phenotypic responses, but higher amounts induced little additional induction. This suggests that additional increases in predator‐induced traits with greater predator risk offer minimal increases in fitness or that a limit in the response magnitude was reached. Additionally, the response thresholds were contingent on the predator and focal trait. For shell width, responses were generally detected at a lower amount of consumed biomass by water bugs compared to crayfish. Within the crayfish treatments, we found that the shell thickness response threshold was lower than the shell width response threshold. When we combined gradients of consumed biomass from both predators, we found that the magnitude of response to one predator was often reduced when the other predator was present. Interestingly, these effects were often detected at consumed biomass levels that were lower than the threshold concentration necessary to elicit a response in the single‐predator treatments. Moreover, our combined predator treatments revealed that snails shifted from discrete responses to more continuous (i.e. graded) responses. Together, our results reveal that organisms experiencing multiple environmental gradients can integrate this information to make phenotypic decisions and demonstrate the novel result that an exposure to multiple species of predators can lower the response threshold of prey.     

Experimental evidence for alternative stable equilibria in a benthic pond food web

Both local and initial conditions have been implicated in causing variation in the strengths of predator effects among natural communities. In this study, I performed a controlled mesocosm experiment using two common snails, Physella gyrina and Helisoma trivolvis, and their insect predator, Belostoma flumineum, to explicitly quantify and understand the causes of previously observed variation in food web interactions in natural ecosystems. The results from a controlled mesocosm experiment showed that: (1) at low productivity, snails were dominated by small‐vulnerable species (P. gyrina), predator effects were strong and predator effects caused a trophic cascade extending to producers; (2) at high productivity, snails were dominated by large‐invulnerable species (H. trivolvis), predator effects were weak and there was no trophic cascade to producers; (3) at intermediate productivity, alternative stable equilibria were implicated, such that depending on initial conditions, snails were dominated by either small‐vulnerable or large‐invulnerable species and predator effects were either strong or weak. Thus, this study represents one of the first to quantify how variation in food web structure can result from the local environment, initial conditions and their interaction.     

Predator-prey relationships among larval dragonflies,salamanders, and frogs

Summary Tadpoles of the barking tree frog, Hyla gratiosa, are abundant in spring and summer in some ponds and Carolina bays on the Savannah River Plant near Aiken, South Carolina. To determine how these tadpoles survive in the presence of predaceous salamander larvae, Ambystoma talpoideum, and larvae of an aeshnid dragonfly, Anax junius, we determined fields densities and sizes of the predators and the prey and conducted predation experiments in the laboratory. Tadpoles rapidly grow to a size not captured by Ambystoma, although Anax larvae can capture slightly larger tadpoles. Differing habitat preferences among the tadpoles and the two predator species probably aid in reducing predation pressure. Preliminary work indicates that the tadpoles may have an immobility response to an attack by a predator. In addition, the smallest, most vulnerable tadpoles have a distinctive color pattern which may function to disrupt the body outline and make them indiscernable to predators.     

How did this snail get here? Several dispersal vectors inferred for an aquatic invasive species

1. How species reach and persist in isolated habitats remains an open question in many cases, especially for rapidly spreading invasive species. This is particularly true for temporary freshwater ponds, which can be remote and may dry out annually, but may still harbour high biodiversity. Persistence in such habitats depends on recurrent colonisation or species survival capacity, and ponds therefore provide an ideal system to investigate dispersal and connectivity. 2. Here, we test the hypothesis that the wide distributions and invasive potential of aquatic snails is due to their ability to exploit several dispersal vectors in different landscapes. We explored the population structure of Physa acuta (recent synonyms: Haitia acuta, Physella acuta, Pulmonata: Gastropoda), an invasive aquatic snail originating from North America, but established in temporary ponds in Doñana National Park, southern Spain. In this area, snails face land barriers when attempting to colonise other suitable habitat. 3. Genetic analyses using six microsatellite loci from 271 snails in 21 sites indicated that (i) geographically and hydrologically isolated snail populations in the park were genetically similar to a large snail population in rice fields more than 15 km away; (ii) these isolated ponds showed an isolation‐by‐distance pattern. This pattern broke down, however, for those ponds visited frequently by large mammals such as cattle, deer and wild boar; (iii) snail populations were panmictic in flooded and hydrologically connected rice fields. 4. These results support the notion that aquatic snails disperse readily by direct water connections in the flooded rice fields, can be carried by waterbirds flying between the rice fields and the park and may disperse between ponds within the park by attaching to large mammals. 5. The potential for aquatic snails such as Physa acuta to exploit several dispersal vectors may contribute to their wide distribution on various continents and their success as invasive species. We suggest that the interaction between different dispersal vectors, their relation to specific habitats and consequences at different geographic scales should be considered both when attempting to control invasive freshwater species and when protecting endangered species.     

Size structure and substitutability in an odonate intraguild predation system

Interactions between different size classes of predator species have the potential to influence survival of prey species in intraguild predation (IGP) systems, but few studies test for these effects. Using a substitutive design in a field setting, I measured the effects of two size classes of IG predators (large and small larvae of the dragonfly Anax junius) on the mortality of IG prey (larvae of the dragonfly Pachydiplax longipennis). I also examined whether combinations of large A. junius and P. longipennis and small A. junius and P. longipennis had substitutable effects on shared prey (larvae of the damselfly Ischnura verticalis). The presence of both size classes of A. junius, when alone and in combination with P. longipennis, significantly increased mortality of I. verticalis. In the presence of P. longipennis, large and small A. junius had similar effects on the mortality of I. verticalis, and effects of size-structured assemblages of A. junius were similar to the effects of each size class alone at the same density. The effects of the two size classes of A. junius on P. longipennis differed, and P. longipennis mortality was lower when exposed to size structured assemblages of A. junius than when exposed to only large A. junius at the same density. Results were similar to those in a laboratory study, although the effect of P. longipennis on I. verticalis was much lower in the field setting. These results demonstrate that interactions between different size classes of IG predators promote the survival of IG prey and highlight the importance of within-species size structure as a characteristic that may promote the coexistence of predators in IGP systems.     

Evaluating the role of regional and local processes in structuring a larval trematode metacommunity of Helisoma trivolvis

Metacommunity theory has advanced our understanding of how local and regional processes affect the structure of ecological communities. While parasites have largely been omitted from metacommunity research, parasite communities can provide the large sample sizes and discrete boundaries often required for evaluating metacommunity patterns. Here, we used assemblages of flatworm parasites that infect freshwater snails (Helisoma trivolvis) to evaluate three questions: 1) what factors affect individual host infections within ponds? 2) Is the parasite metacommunity structured among ponds? And 3) what is the relative role of local versus regional processes in determining metacommunity structure and species richness among ponds? We examined 10 821 snails from 96 sites in five park complexes in the San Francisco Bay area, California, and found 953 infections from six parasite groups. At the within‐pond level, infection status of host snails correlated positively with individual snail size and pond infection prevalence for all six parasite groups. Using an ordination method to test for metacommunity structure, we found that the parasite metacommunity was organized in a non‐random pattern with species responding individually along an environmental gradient. Based on a model selection approach involving local and regional predictors, parasite species richness and metacommunity structure correlated with both local abiotic (pH and total dissolved nitrogen) and biotic (non‐host mollusk density, and H. trivolvis biomass) factors, with little support for regional predictors. Overall, this trematode metacommunity most closely followed the predictions from the species sorting or mass effects metacommunity paradigm, in which community diversity is filtered by local site characteristics.     

Foraging and refuge use by a pond snail: Effects of physiological state,predators, and resources

The costs and benefits of anti-predator behavioral responses should be functions of the actual risk of predation, the availability of the prey's resources, and the physiological state of the prey. For example, a food-stressed individual risks starvation when hiding from predators, while a well-fed organism can better afford to hide (and pay the cost of not foraging). Similarly, the benefits of resource acquisition are probably highest for the prey in the poorest state, while there may be diminishing returns for prey nearing satiation. Empirical studies of state-dependent behavior are only beginning, however, and few studies have investigated interactions between all three potentially important factors. Here I present the results of a laboratory experiment where I manipulated the physiological state of pond snails (Physa gyrina), the abundance of algal resources, and predation cues (Belostoma flumineum waterbugs consuming snails) in a full factorial design to assess their direct effects on snail behavior and indirect effects on algal biomass. On average, snails foraged more when resources were abundant, and when predators were absent. Snails also foraged more when previously exposed to physiological stress. Snails spent more time at the water's surface (a refuging behavior) in the presence of predation cues on average, but predation, resource levels, and prey state had interactive effects on refuge use. There was a consistent positive trait-mediated indirect effect of predators on algal biomass, across all resource levels and prey states.     

Lethal and non-lethal effects of multiple indigenous predators on the invasive golden apple snail (Pomacea canaliculata)

1. We investigated the individual and combined effects of two predators (the climbing perch, Anabas testudineus, and the wetland crab, Esanthelphusa nimoafi) indigenous to wetlands in Laos, on the behaviour and survival of the invasive South American golden apple snail (Pomacea canaliculata). The snail is considered a pest, consuming large amounts of rice and other aquatic vegetation in the region. 2. Snail avoidance reactions to released predator chemical cues were investigated in aquaria while the effects of predators on a mixed snail population were studied in field enclosures that contained native aquatic plants (Salvinia cucullata, Ludwigia adscendens and Ipomoea aquatica). 3. In the aquaria experiment, neonate (2–3 mm) and medium‐sized snails (8–10 mm) responded to fish chemical cues by going to the surface, whereas adult snails (35–40 mm) went to the bottom. In contrast, no size class of snails reacted to chemical cues released by crabs. 4. In the field experiment, fish reduced the abundance of neonate snails, and crabs reduced the abundance of all size classes. The effect of the combined predators could not be predicted from the mortality rate observed in single predator treatments. The survival of neonate and medium‐sized snails was greater and of adults less than expected. The presence of predators did not affect egg production. Snails consumed significant amounts of plants despite the presence of predators. 5. Our findings suggest that some indigenous Asian predators have lethal and sublethal effects on P. canaliculata that depend on snail size and predator type. When in the presence of several predators the response of snails to one predator may either increase or decrease the vulnerability of snails to the others.     

Observations of insect predation on rotifers

Interactions between rotifers and their insect predators have not received adequate attention, possibly due to the assumption that rotifers are too small for insects to eat. In laboratory experiments, we offered the rotifers Hexarthra mira, Plationus patulus and small and large Synchaeta pectinata to four common insect predators: the notonectids Notonecta lunata and Buenoa macrotibialis, the smaller hemipteran Neoplea striola and small (1.5 mm) aeschnid dragonfly larvae. Excepting Plationus offered to dragonflies, all rotifer preys were consumed to some degree. No size selectivity was apparent for predators that ate few rotifers, but small instar Buenoa ate significantly more large (420 m) than small (300 m) Synchaeta. Predator size appeared to be less important than predatory style and prey morphology in determining ingestion rates. Neoplea and dragonflies ate more Hexarthra than Plationus, while the pattern was reversed for Buenoa, possibly because Buenoa is able to manipulate the hard lorica of Plationus better. Insect predators are capable of direct suppression of rotifer populations, an interaction which may be particularly important in littoral zones and fishless ponds where macroinvertebrates are numerous.     

Habitat selection in the larvae of two species of Zygoptera (Odonata): biotic interactions and abiotic limitation

Field and laboratory experiments were set up to obtain data on the reasons for different habitat selection of Enallagma cyathigerum and Platycnemis pennipes. (1) Rearing of larvae in two different ponds showed that while P. pennipes was not able to survive conditions of low oxygen content, 50% of the E. cyathigerum larvae survived. (2) In field predation experiments with sticklebacks and dragonflies as predators, we found that E. cyathigerum suffered highest predation by the fish. In P. pennipes, mortality was highest with Anax imperator. (3) Experiments regarding larval behaviour showed that E. cyathigerum was generally more active and had higher foraging success than P. pennipes. Both species reduced activity in the presence of fish, but E. cyathigerum did so to a minor extent. In contrast to P. pennipes, E. cyathigerum showed escaping behaviour. (4) In the laboratory, the growth of E. cyathigerum was faster than that of P. pennipes.     

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.     

Experimental Test of Preferences for an Invasive Prey by an Endangered Predator: Implications for Conservation

Identifying impacts of exotic species on native populations is central to ecology and conservation. Although the effects of exotic predators on native prey have received much attention, the role of exotic prey on native predators is poorly understood. Determining if native predators actively prefer invasive prey over native prey has implications for interpreting invasion impacts, identifying the presence of evolutionary traps, and predator persistence. One of the world’s most invasive species, Pomacea maculata, has recently established in portions of the endangered Everglade snail kite’s (Rostrhamus sociabilis plumbeus) geographic range. Although these exotic snails could provide additional prey resources, they are typically much larger than the native snail, which can lead to lower foraging success and the potential for diminished energetic benefits in comparison to native snails. Nonetheless, snail kites frequently forage on exotic snails. We used choice experiments to evaluate snail kite foraging preference in relation to exotic species and snail size. We found that snail kites do not show a preference for native or exotic snails. Rather, snail kites generally showed a preference for medium-sized snails, the sizes reflective of large native snails. These results suggest that while snail kites frequently forage on exotic snails in the wild, this behavior is likely driven simply by the abundance of exotic snails rather than snail kites preferring exotics. This lack of preference offers insights to hypotheses regarding effects of exotic species, guidance regarding habitat and invasive species management, and illustrates how native-exotic relationships can be misleading in the absence of experimental tests of such interactions.     

Terrestrial snails use predator-diet to assess danger

Some aquatic snails are able to use chemical cues (kairomones) to differentiate between predators that have fed on snails and predators that have eaten other prey. However, it is unknown if terrestrial snails are able to differentiate between snail-fed predators and predators that have not recently consumed snails. Here we document diet-based chemical discrimination of a predator, the ground beetle Carabus carabus, by a terrestrial snail Theba pisana. When exposed to the feces of snail-fed beetles, snails initially stopped all movements and then increased climbing speed. The snails also decreased the time to deposition of their egg clutch. The snails did not react to an extract of crushed snails. Snails had only a partial reaction to the feces of beetles that had fed on chicken (Gallus domesticus) livers—they decreased climbing speed but did not alter egg laying times. These responses may be adaptive in that they allow snails to differentiate between individual beetles that may pose an immediate threat and beetles that may not. This is one of only a few studies to examine predator-diet effects on reproductive behavior.     

The rules of engagement: how to defend against combinations of predators

Studies of inducible defenses have traditionally examined prey responses to one predator at a time. However, prey in nature encounter combinations of predators that should force them to produce phenotypic compromises. We examined how snails (Helisoma trivolvis) alter their phenotype in the presence of three different predator species that were presented alone and in pairwise combinations. When snails were exposed to each predator alone, they formed predator-specific defenses that reflected the differences in each predator’s foraging mode. When snails were exposed to pairwise combinations of predators, their phenotype was dependent on their ability to detect each predator, the risk posed by each predator, and the effectiveness of a given defense against each predator. Consequently, responses to combined predators were typically biased towards one of the predators in the pair. This suggests that prey facing combined predators do not form simple intermediate defenses and, as a result, may experience enhanced mortality risk when they encounter natural predator regimes.     

Wind Drift Compensation in Migrating Dragonflies Pantala (Odonata: Libellulidae)

Tailwind drift compensation serves to maximize a migrant's flight distance on a given amount of energy, and crosswind drift compensation serves to hold a course true and minimize the distance flown. With full or part compensation, airspeeds are predicted to increase with greater crosswind drift. To test whether migrating dragonflies compensated for wind drift, I measured the velocity and heading of Pantala hymenaea and P. flavescens in natural flight over a lake and the ambient wind speed and direction. P. hymenaea flew north-easterly (58°), whereas P. flavescens flew significantly more east–north easterly (74°) throughout the day. Pantala spp. demonstrated part compensation for changes in crosswind drift within individuals (mean compensation = 54%, P = 0.0000), evidence for use of a ground reference to correct for drift when flying over water. Among individuals, P. flavescens compensated for crosswind drift. P. hymenaea overcompensated and then drifted downwind on one morning and compensated for crosswind drift on the next. As predicted from optimal migration theory, airspeed (5.0 m/s for both species with no tailwind) decreased with tailwind velocity both among individuals (data for both species pooled [n = 19], P < 0.0001) and within each individual as it crossed the lake (P = 0.0016).     

Indirect effects of a parasite on a benthic community: an experiment with trematodes, snails and periphyton

1. Few studies have directly addressed the role played by parasites in the structure and function of ecosystems. Parasites influence the behaviour, reproduction and overall fitness of their hosts, but have been usually overlooked in community and ecosystem‐level studies. We investigated the effects of trematode parasites on snail–periphyton interactions. 2. Physa  acuta (Gastropoda: Pulmonata) snails infected with the trematode Posthodiplostomum minimum (often >30% of within‐shell biomass) grazed more rapidly than uninfected snails. Trematode effects on snail grazing indirectly affected the standing stock and community structure of periphyton. Populations of snails with 50% infected individuals reduced algal biomass by 20% more than populations with lesser (10% or 0%) infection rates. 3. The alga Cladophora glomerata dominated periphyton communities grazed by snail populations with 50% infection rates, whereas diatoms and blue–green algal taxa dominated when grazed by snail populations with lower infection rates. 4. Thus, trematodes indirectly affected periphyton communities by altering host snail behaviour, a trait‐mediated indirect effect. These results indicate that trematodes can indirectly influence benthic community structure beyond simple population fitness, with possible related effects on ecosystem function.