Predator identity and consumer behavior: differential effects of fish and crayfish on the habitat use of a freshwater snail

Predators can alter the outcome of ecological interactions among other members of the food web through their effects on prey behavior. While it is well known that animals often alter their behavior with the imposition of predation risk, we know less about how other features of predators may affect prey behavior. For example, relatively few studies have addressed the effects of predator identity on prey behavior, but such knowledge is crucial to understanding food web interactions. This study contrasts the behavioral responses of the freshwater snail Physellagyrina to fish and crayfish predators. Snails were placed in experimental mesocosms containing caged fish and crayfish, so the only communication between experimental snails and their predators was via non-visual cues. The caged fish and crayfish were fed an equal number of snails, thereby simulating equal prey mortality rates. In the presence of fish, the experimental snails moved under cover, which confers safety from fish predators. However, in the presence of crayfish, snails avoided benthic cover and moved to the water surface. Thus, two species of predators, exerting the same level of mortality on prey, induced very different behavioral responses. We predict that these contrasting behavioral responses to predation risk have important consequences for the interactions between snails and their periphyton resources. Received: 1 June 1998 / Accepted: 12 October 1998     

Chemical cues modify species interactions: the ecological consequences of predator avoidance by freshwater snails

Chemical signals released by predators or injured prey often induce shifts in the traits of prey species, which may in turn affect species interactions. Here we investigate the role that chemical cues play in mediating species interactions in the littoral food web of lakes. Previous studies have shown that predators induce shifts in the morphology, life history, and behavior of the freshwater snail Physella, but the ecological consequences of developing these inducible defenses are not well documented. We observed habitat use of the freshwater snail Physella gyrina along a depth gradient in a natural lake, and found they increased their use of covered habitats with increasing depth. We hypothesized that this habitat shift was due to changes in the level and type of predation risk, and that the habitat shift would affect periphyton standing crops. These hypotheses were tested in a mesocosm experiment in which we manipulated the presence of molluscivorous fish and crayfish. Predators were confined to cages and snail density was identical in all treatments, so any effects of predators were mediated through trait shifts induced by chemical cues. In the presence of fish, Physella moved under cover, but in the presence of crayfish, Physella avoided cover and moved to the water surface. These non‐lethal effects of predators on snail habitat use influenced the interaction between snails and their periphyton resources. In the presence of fish, periphyton standing crop in covered habitats was reduced to just 8% of periphyton in the absence of fish. Crayfish had no significant effect on periphyton in covered habitats, but they reduced periphyton in near‐surface habitats to 39% of the standing crop in the absence of crayfish. The combined effects of fish and crayfish were generally intermediate to their individual effects. We conclude that because chemical cues often have strong effects on individual traits and trophic interactions are sensitive to trait values, chemical cues may play an important role in shaping the structure and dynamics of food webs.     

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

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

Non-lethal effects of predators on prey growth rates depend on prey density and nutrient additions

A number of studies show that predators can depress prey growth rates by inducing reductions in foraging activity, but the size of this non-lethal effect is quite variable. Here I investigate how prey density and resource productivity may alter the extent to which predators depress the growth rates of their prey. Theory predicts that when resources are overgrazed, an increase in predation risk will have little net effect on individual food intake because the decline in foraging effort will be offset by an increase in resource level. Thus, the non-lethal effects of predators on prey growth rates should depend upon prey density and resource productivity in a predictable manner, with the growth penalty imposed by predators being strongest when resources are undergrazed and weakest when resources are overgrazed. I tested this hypothesis by manipulating predation risk, prey density, and nutrient additions in a mesocosm experiment with the pulmonate snail Helisoma trivolvis . Refuge use by snails was 45% higher in the presence of caged crayfish than in their absence. Snail growth rates were reduced, on average, by 24% in the presence of caged crayfish. However, the magnitude of the growth penalty exacted by crayfish depended on snail density and nutrient additions. When snails were stocked at high density and nutrient additions were low, growth suppression was just 2.6%. At the other extreme, when snails were at low density and nutrient additions were high, growth suppression was 44.6%. Thus, the non-lethal effects of predators on prey growth depend on environmental context, illustrating an important link between individual traits and system-level properties.     

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.     

Contrasting effects of an invasive crayfish (Procambarus clarkii) on two temperate stream communities

1. The effects of omnivorous exotic species on native communities are often difficult to predict because of the broad diets and behavioural flexibility of the omnivore, and the diverse abiotic and biotic characteristics of invaded systems. We investigated experimentally the effects of a gradient of density of the introduced, omnivorous red swamp crayfish Procambarus clarkii (Decapoda: Cambaridae) on two stream communities in southern California, U.S.A. 2. The Ventura River is a clear, flowing stream with a cobble substratum, with abundant algae but low densities of large invertebrates, small herbivores and snails. The Santa Ynez River at the time of the study consisted of a series of drying pools underlain by sand, with abundant charophytes, large predatory invertebrates and herbivores, including snails. 3. In the Ventura River, periphyton biomass and inorganic sediment decreased with increasing crayfish abundance, but in the Santa Ynez River, periphyton and sediment were unrelated to crayfish densities. 4. In the Ventura River, the biomass and density of all benthic invertebrates combined, chironomids, micropredators, the meiofauna (chydorid cladocerans, copepods and ostracods), and specific predatory and herbivorous taxa, as well as taxon richness, were negatively related to crayfish density. In the Santa Ynez River, the biomass and average body size of benthic invertebrates, predatory invertebrates, herbivores and chironomids, but not total invertebrate density or taxon richness, were negatively related to crayfish density. 5. Fewer large predatory invertebrates and snails (Physella gyrina) in both streams, and baetid mayflies in the Ventura River, were visible at night in channels where crayfish were abundant. Snails responded to crayfish by moving above the water line in the Santa Ynez River, but not in the Ventura River. 6. We suggest that the same omnivore had different effects on these neighbouring streams because of crayfish predation on large invertebrates in the Santa Ynez River and the scarcity of such prey in the Ventura River, leading to increased crayfish grazing on periphyton, and reductions in periphyton‐associated invertebrates, in the Ventura River.     

Home-field advantage: native signal crayfish (Pacifastacus leniusculus) out consume newly introduced crayfishes for invasive Chinese mystery snail (Bellamya chinensis)

The introduction of non-indigenous plants, animals and pathogens is one of today’s most pressing environmental challenges. Freshwater ecologists are challenged to predict the potential consequences of species invasions because many ecosystems increasingly support novel assemblages of native and non-native species that are likely to interact in complex ways. In this study we evaluated how native signal crayfish (Pacifastacus leniusculus) and non-native red swamp crayfish (Procambarus clarkii) and northern crayfish (Orconectes virilis) utilize a novel prey resource: the non-native Chinese mystery snail (Bellamya chinensis). All species are widespread in the United States, as well as globally, and recent surveys have discovered them co-occurring in lakes of Washington State. A series of mesocosm experiments revealed that crayfish are able to consume B. chinensis, despite the snail’s large size, thick outer shell and trapdoor defense behaviour. Crayfish exhibited size-selective predation whereby consumption levels decreased with increasing snail size; a common pattern among decapod predators. Comparison of prey profitability curves—defined as the yield of food (weight of snail tissue) per second of feeding time (the time taken to crack the shell and consume the contents)—suggests that small and very large snails may represent the most profitable prey choice. By contrast, previous studies have reported the opposite pattern for crayfish consumption on thin-shelled snails. For all snail size classes, we found that native P. leniusculus and invasive O. virilis consumed greater numbers of snails than invasive P. clarkii. Moreover, P. leniusculus consistently handled and consumed snails at a faster pace compared to both invasive crayfishes across the range of snail sizes examined in our study. These results suggest not only that B. chinensis is a suitable food source for crayfish, but also that native P. leniusculus may ultimately out-consume invasive crayfishes for this new prey resource.     

The influence of temporally variable predation risk on indirect interactions in an aquatic food chain

We know little about how temporally variable predation risk influences prey behavior. The risk allocation hypothesis predicts that prey facing more frequent risk should show weak anti-predator responses, and should be particularly active foragers during rare periods of safety, compared to prey facing infrequent risk. Several studies offer support for the risk allocation hypothesis, but how these responses might propagate through the larger ecological community remains largely unknown. We experimentally investigated the relative strength of trait- and density-mediated indirect effects of a predator on its prey’s resource across predation treatments that varied the lethality (caged or free-swimming predators) and temporal variability (always, often, or sometimes present) of predation. We performed this experiment in pond mesocosms using a giant water bug predator (Belostoma lutarium), an herbivorous pond snail (Physa gyrina), and algae as the basal resource. Snails greatly reduced the abundance of their algal resource when in the absence of predation. Lethal predation at low and medium intensities had significant positive indirect effects on the abundance of algae, mostly by reducing snail density. Snails responded behaviorally to high levels of deadly predation by foraging more and hiding less than in other situations, as predicted by the risk allocation hypothesis, and thus ameliorated the density-mediated indirect effects of predators on algae. Behavioral responses to caged predators, and the subsequent trait-mediated indirect effects, were negligible regardless of predation intensity. Our previous work has demonstrated that trait-mediated indirect effects are weak when resources are abundant, as they were in this experiment. This work demonstrates that temporal variation in predation intensity plays a key role in determining the relative strength of TMIIs and DMIIs in an aquatic food chain.     

Dragonfly predators influence biomass and density of pond snails

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.     

Temporal environmental variation and phenotypic plasticity: a mechanism underlying priority effects

Understanding the role of history in the formation of communities has been a major challenge in community ecology. Here, we explore the role of phenotypic plasticity and its associated trait‐mediated indirect interactions as a mechanism behind priority effects. Using organisms with inducible defenses as a model system, we examine how aquatic communities initially containing different predator environments are affected at the individual and community level by the colonization of a second predator. Snails and tadpoles were established in four different caged‐predator environments (no predator, fish, crayfish or water bugs). These four communities were then crossed with three predator colonization treatments (no colonization, early colonization, or late colonization) using lethal water bugs as the predator. The snails responded to the caged predator environments with predator‐specific behavioral and morphological defenses. In the colonization treatments, snails possessing the wrong phenotype attempted to induce phenotypic changes to defend themselves against the new risk. However, snails initially induced by a different predator environment often suffered high predation rates. Hence, temporal variation in predation risk not only challenged the snail prey to try to track this environmental variation through time by adjusting their defensive phenotypes, but also caused trait‐mediated interactions between snails and the colonizing predator. For tadpoles within these communities, there was little evidence that the morphological responses of snails indirectly effected tadpole predation rates by colonizing water bugs. Unexpectedly, predation rates on tadpoles by colonizing water bugs were generally higher in the three caged‐predator treatments, suggesting that water bugs elevated their foraging activity in response to potentially competing predators. In summary, we demonstrate an important priority effect in which the initial occurrence of one species of predator can facilitate predation by a second predator that colonizes at a later date (i.e. a TMII) suggesting that phenotypic plasticity can be an important driver behind priority effects (i.e. historical exposure to predators).     

Predatory Potential of Freshwater Animals on an Invasive Agricultural Pest, the Apple Snail Pomacea canaliculata (Gastropoda: Ampullariidae), in Southern Japan

The apple snail Pomacea canaliculata is an invasive species and a serious pest of rice in many Asian countries. We studied predatory activities of various animals living in Japanese freshwater habitats, by keeping each individual of a potential predator species with 36 snails of various sizes for three days in the aquarium. Forty-six species were tested, and 26 in eight classes fed on small snails. A species of leech, crabs, the common carp, turtles, the mallard duck and the Norway rat attacked even adult snails of 20–30 mm in shell height. These findings will be helpful in identifying effective predators for biological control of the pest snail. In addition, most of the animals attacking snails are reported to be common in rivers or ponds, but few live in modernized paddy fields having little connections with natural water systems. This may be a reason why this snail maintains large populations in paddy fields but not in other freshwater habitats.     

Location of Biomphalaria pfeifferi by Schistosoma mansoni miracidia in stagnant and running water under field conditions

The efficiency of S. mansoni miracidia in locating and infecting Biomphalaria pfeifferi in Gezira canals has been studied under field conditions. When S. mansoni eggs were introduced into clean stagnant water in small field channels, the miracidia hatched to infect 100% of 30 snails in cages at the release point. Fifteen metres upstream and downstream 13% of caged snails were infected but no infections were found in snails 20 m away.When eggs were released into the same canal in flowing water (8.3 cm · s^–1), no infections were detected in any of the caged snails placed 0–100 m downstream. Releasing hatched miracidia instead of eggs resulted in infections in all cages at 5 m intervals from 0-100 m. The release of eggs into flowing water was likened to the method by which S. haematobium eggs are deposited during urination. The 0% infection suggests that eggs will be swept away from the point of contamination by the flow. Thus only urination into stagnant water will lead to heavy snail infection rates.When eggs were released into a small pond-like minor canal tail end snail infection rates were only 3%. This was probably due to the larger water volume, smaller number of caged snails, and the presence of vegetation and other fauna which may be decoys or predators.The results highlight how very high snail infection rates can be produced under ideal conditions but also show how large snail and miracidia numbers are required in natural situations.     

Cooccurrence of prey species alters the impact of predators on prey performance through multiple mechanisms

When prey are differentially affected by intra and interspecific competition, the cooccurrence of multiple prey species alters the per capita availability of food for a particular prey species which could alter how prey respond to the threat of predation, and hence the overall‐effect of predators. We conducted an experiment to examine the extent to which the nonconsumptive and overall effect of predatory water bugs on snail and tadpole traits (performance and morphology) depended on whether tadpoles and snails cooccurred. Tadpoles and snails differed in their relative susceptibility to intraspecific and interspecific competition, and predators affected both prey species via consumptive and nonconsumptive mechanisms. Furthermore, the overall effect of predators often depended on whether another prey species was present. The reasoning for why the overall effect of predators depended on whether prey species cooccurred, however, differed for each of the response variables. Predators affected snail body growth via nonconsumptive mechanisms, but the change in the overall effect of predators on snail body growth was attributable to how snails responded to competition in the absence of predators, rather than a change in how snails responded to the threat of predation. Predators did not affect tadpole body growth via nonconsumptive mechanisms, but the greater vulnerability of competitively superior prey (snails) to predators increased the strength of consumptive mechanisms (and hence the overall effect) through which predators affected tadpole growth. Predators affected tadpole morphology via nonconsumptive mechanisms, but the greater propensity for predators to kill competitively superior prey (snails) enhanced the ability of tadpoles to alter their morphology in response to the threat of predation by creating an environment where tadpoles had a higher per capita supply of food available to invest in the development of morphological defenses. Our work indicates that the mechanisms through which predators affect prey depends on the other members of the community.     

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.     

Fleeing towards death – leech‐induced behavioural defences increase freshwater snail susceptibility to predatory fish

Prey species are often exposed to multiple predators, which presents several difficulties to prey species. This is especially true when the response to one predator influences the prey’s susceptibility to other predators. Predator‐induced defences have evolved in a wide range of prey species, and experiments involving predators with different hunting strategies allow researchers to evaluate how prey respond to multiple threats. Freshwater snails are known to respond to a variety of predators with both morphological and behavioural defences. Here we studied how freshwater snails Radix balthica responded behaviourally to fish and leech predators, both separately and together. Our aim was to explore whether conflicting predator‐induced responses existed and, if so, what effect they had on snail survival when both predatory fish and leeches were present. We found that although R. balthica increased refuge use when exposed to predatory fish, they decreased refuge use when exposed to predatory leeches. When both predators were present, snails showed a stronger response towards leech than fish and responded by leaving the refuge. This response made the snails more susceptible to fish predation, which increased snail mortality when exposed to both fish and leech compared to fish only. We show that predators that have a relatively low predation rate can substantially increase mortality rates by indirect effects. By forcing snails out of refuges such as rock and macrophyte habitats, leeches can indirectly increase predation from molluscivorous fish and may thus affect snail densities.     

The long‐term impacts of predators on prey: inducible defenses,population dynamics,and indirect effects

Despite the amount of research on the inducible defenses of prey against predators, our understanding of the long‐term significance of non‐lethal predators on prey phenotypes, prey population dynamics, and community structure has rarely been explored. Our objectives were to assess the effects of predators on prey defenses, prey population dynamics, and the relative magnitude of density‐ versus trait‐mediated indirect interactions (DMIIs and TMIIs) over multiple prey generations. Using a freshwater snail and three common snail predators, we constructed a series of community treatments with pond mesocosms that manipulated trophic structure, the identity of the top predator, and whether predators were caged or uncaged. We quantified snail phenotypes, snail population size, and resource abundance over multiple snail generations. We found that snails were expressing inducible defenses in our system although the magnitude of the responses varied over time and across predator species. Despite the expression of inducible defenses, caged predators did not reduce snail population size. There also was no evidence of TMIIs throughout the experiment suggesting that TMIIs have a minimal role in the long‐term structure of our communities. The absence of TMIIs was largely driven by the lack of predator‐induced reductions in resource consumption and the lack of consistent reductions in population size with predator cues. In contrast, we detected strong DMIIs associated with lethal predators suggesting that DMIIs are the dominant long‐term mechanism influencing community structure. Our results demonstrate that although predators can have significant effects on prey phenotypes and sometimes cause short‐term TMIIs, there may be few long‐term consequences of these responses on population dynamics and indirect interactions, at least within simple food webs. Research directed towards addressing the long‐term consequences of predator–prey interactions within communities will help to reveal whether the conclusions and predictions generated from short‐term experiments are applicable over ecological and evolutionary timescales.     

The potential for using red claw crayfish and hybrid African catfish as biological control agents for Schistosoma host snails

The potential of red claw crayfish and hybrid African catfish (Clarias gariepinus and Clarias ngamensis) as predators for Schistosoma host snails was evaluated in 2014 by monitoring the consumption of snails by crayfish and catfish in experimental tanks over time under laboratory conditions. After 15 days, both crayfish and catfish had significantly reduced the populations of Bulinus globosus. Crayfish consumed 6.9 snails d^?1, whereas catfish consumed 5.9 snails d^?1. However, when supplied with an alternative prey, Melanoides tuberculata (100 individuals per tank), crayfish clearly preferred M. tuberculata (100% consumed over seven days) to B. globosus (100 individuals per tank) (54% consumed over the same period). Catfish, conversely, did not have a clear preference for either prey species, consuming 77% and 88% of M. tuberculata and B. globosus, respectively. It was also observed that young catfish were more efficient predators than older ones, because of ontogenetic shifts in their diet with age. Hybrid catfish retain the molluscivorous characteristics of their parent stock and red claw crayfish also preys on Schistosoma host snails. However, the effectiveness of both predators is affected by the presence of an alternative prey. Therefore, under suitable conditions, these species can be considered for biological control of schistosomiasis transmission.     

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.     

Multiple predator-avoidance behaviours of the freshwater snail Physella heterostropha pomila: responses vary with risk

• 1 We examined the predator‐avoidance behaviour, exhibited in response to chemical cues, of two populations of the snail Physella heterostropha pomila. Snails were subjected to four treatments simulating different degrees of predation risk: control water (low risk), or water from tanks containing nonforaging crayfish (intermediate risk), crushed conspecifics (high risk) or crayfish consuming conspecifics (high risk). Data were analysed using three‐way ANOVA models (population × predator chemicals × injured conspecific chemicals).
• 2 Physella increased its avoidance behaviour as risk increased. Crayfish cue elicited a significantly greater response than from controls. Cues from injured conspecifics elicited the strongest response.
• 3 Physella exhibited several types of avoidance behaviour, including burial into the substratum, moving to the water surface, and crawling out of the water. The type of cue present influenced response type. Cues from crayfish reduced burial and increased movement to the water surface or out of the water. Cues from injured‐conspecifics significantly increased crawling completely out of the water.
• 4 The two populations differed in the type and degree of response exhibited. One population exhibited significantly greater ‘reactivity’ (i.e. any avoidance behaviour) in response to foraging crayfish, and more burial and crawl‐out behaviours were exhibited in high‐risk treatments.

     

Life-history strategies of a freshwater snail in response to stream permanence and predation: balancing conflicting demands

Summary Life-history parameters for the freshwater snail Physella virgata virgata were estimated in temporary and permanent streams with and without crayfish (known snail predators). Snails from the permanent stream with crayfish exhibited higher age and size at first reproduction, as well as higher size and age at death, compared to snail populations from both temporary streams and the permanent stream without crayfish. Life-history patterns of snails from the permanent and temporary streams without crayfish were indistinguishable, with small sizes at first reproduction and at death, as well as lower age at first reproduction and at death. The snail population from the temporary stream with crayfish showed an intermediate life-history pattern. The results suggest that snail life-history patterns vary with both predation pressure and habitat stability, exhibiting phenotypically plastic responses to the variables.