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

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

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.     

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

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

Biological control of invasive apple snails by two species of carp: Effects on non-target species matter

Molluscivorous fish, especially carp, have been adopted as bio-control agents of the invasive apple snail Pomacea canaliculata, but previous studies have focused on their effectiveness, with little attention paid to their undesirable effects on non-target plants and animals. We conducted an 8-week mesocosm study to compare the effectiveness of two indigenous fish, common carp (Cyprinus carpio) and black carp (Mylopharyngodon piceus), in removing P. canaliculata, and their potential side effects on macrophytes and non-target mollusks in a freshwater wetland. Three species of macrophytes and a community of mollusks in the wetland sediment were enclosed in 1 × 1 × 1 m enclosures either with apple snails (AS), with apple snails and common carp (AS + CC), with apple snails and black carp (AS + BC), or without apple snails and fish. Both species of carp were effective predators of P. canaliculata, removing most of the individuals in the enclosures except a few that were too big to fit into their mouth. By reducing apple snail population, black carp reduced grazing of apple snail on macrophytes. In contrast, although common carp controlled apple snail population, it did not reduce overall loss in plant biomass as the fish might also fed on macrophytes. Both species of carp preyed on non-target mollusks. Application of bio-control agents in invasive species management needs to consider their effects on both the pest and non-target plants and animals. Adoption of common and/or black carp to control apple snail populations thus depends on the weight given to their effectiveness and subtle different effects on non-target organisms by wetland management authority.     

Size-dependent effects of an invasive herbivorous snail (Pomacea canaliculata) on macrophytes and periphyton in Asian wetlands

1. The invasive golden apple snail (Pomacea canaliculata), native to South America, is a serious pest on rice seedlings in south‐east Asia and has also been shown to consume large amounts of macrophytes in natural wetlands, with large effects on ecosystem functioning. Earlier studies suggest that the snail undergoes an ontogenetic diet shift, feeding on algae and detritus as juveniles and shifting to aquatic macrophytes as adults. 2. Here, we study the effects of snail populations with a size‐structure typical of either populations at an invasive front or the size‐structure of established populations. In an enclosure experiment performed in a wetland in Laos, we compared treatments with small snails only (3 mm; invasive treatment) to treatments with small, medium sized (10 mm) and adult (>25 mm) snails (established treatment). The effects of snail grazing on three aquatic macrophyte species and periphytic algae were quantified. 3. We found that snails of all sizes had a strong negative effect on the biomass of all macrophyte species and periphytic algae. There was no evidence of an ontogenetic diet change, i.e. snails in both the invasive and established treatments affected macrophyte biomass. Foraging was size‐dependent in that small snails had higher relative foraging capacity (g plant consumed per g of snail) compared with medium and adult snails. Small snails, therefore, depressed growth of medium snails at increasing densities through exploitative competition for preferred resources, while adult snails did not grow at all in the presence of small snails. 4. Density dependence is common in freshwater invertebrates, including gastropod populations, but differences in size dependent foraging‐ and competitive‐ability have rarely been demonstrated in this group of organisms. Knowledge about intra‐specific differences in ecological performance may, however, both deepen our understanding of the processes that underlie population dynamics in invertebrates such as gastropods, and help develop control strategies for invasive golden apple snails.     

Complex interactions among fish, snails and macrophytes: implications for biological control of an invasive snail

The golden apple snail (Pomacea canaliculata), a native of freshwater wetlands of South America, has invaded many Asian countries and grazed heavily in agricultural and wild areas. Common carp (Cyprinus carpio) has been proposed as a biological control agent against this snail, but little is known about its impact on non-target aquatic plants and animals. In a 8-week enclosure experiment, we quantified the impact of common carp on three species of aquatic macrophytes and nine species of snails, including the apple snail, in a shallow pond. The results showed that the apple snail or carp alone significantly reduced the plant biomass, although the apple snail had a stronger overall herbivorous effect than the carp. The carp completely removed juvenile apple snails, but had only a weak predatory effect on larger apple snails and no effect on the adults’ oviposition frequency. Furthermore, the carp significantly reduced the populations of most species of other snails that occurred naturally in the pond. Our results thus indicate that common carp can be an effective biological control agent against the invasive apple snail, but caution should be taken about its potential to reduce wetland floral and faunal diversity.     

Assessing density–damage relationships between water hyacinth and its grasshopper herbivore

Plants are variable in their responses to insect herbivory. Experimental increases in densities of phytophagous insects can reveal the type of plant response to herbivory in terms of impact and compensatory ability. The relationship between insect density and plant damage of a grasshopper, Cornops aquaticum Brüner (Orthoptera: Acrididae: Tetrataeniini), a candidate biological control agent, and an invasive aquatic plant, water hyacinth, Eichhornia crassipes Mart. Solms‐Laubach (Pontederiaceae), was investigated to assess potential damage to the weed. The impact of different densities of male and female grasshoppers on E. crassipes growth parameters was determined in a quarantine glasshouse experiment. Damage curves indicated that the relationship between plant biomass reduction and insect density was curvilinear whereas leaf production was linear. Female C. aquaticum were more damaging than males, causing high rates of plant mortality before the end of the trial at densities of three and four per plant. Feeding by C. aquaticum significantly reduced the total plant biomass and the number of leaves produced, and female grasshoppers caused a greater reduction in the number of leaves produced by water hyacinth plants than males. Grasshopper herbivory suppressed vegetative reproduction in E. crassipes, suggesting C. aquaticum could contribute to a reduction in the density and spread of E. crassipes infestations. The results showed that E. crassipes vigour and productivity decreases with an increase in feeding intensity by the grasshopper. Cornops aquaticum should therefore be considered for release in South Africa based on its host specificity and potential impact on E. crassipes.     

Snails prefer it sweet: A multifactorial test of the metal defence hypothesis

Metal defence against insect herbivory in hyperaccumulator plants is well documented. However, there are contradictory results regarding protection against snails. According to the joint effects hypothesis, inorganic and organic defences cooperate in plant protection. To test this hypothesis, we explored the relationships between snail (Cantareus aspersus) feeding and multiple inorganic and organic leaf components in the Cd hyperaccumulator plant Noccaea praecox. Plants grouped by rosette size growing in nutrient solution supplemented or not with 50 μM Cd were offered to the snails. After 3 days of snail feeding, the plants and snails were analysed. In addition to Cd concentrations, we analysed leaves for nutritional factors (sugar and protein), defence‐related compounds (glucosinolates, phenolics, tannins, salicylic acid and jasmonate) and essential mineral nutrients. Cadmium concentrations in the snails and in snail excrements were also analysed. Snails preferentially fed on plants grown without Cd. Medium‐sized plants exposed to Cd were the least consumed. Snail excrements from this trial weighed less and had higher Cd concentrations than those from other treatments. Cadmium increased salicylate and jasmonate production. A positive relationship between jasmonate levels and the number of attacked leaves was found. Principal component analysis revealed that leaf sugar concentration was the main factor positively affecting snails' leaf consumption, while leaf Cd had a negative but weaker influence. In conclusion, leaf sugar concentration mainly governs snails' feeding preferences. High leaf Cd concentrations do not deter herbivores from attacking leaves, but they do reduce leaf consumption. Our results clearly support the joint effects hypothesis.     

Indirect interactions in a rice ecosystem: density dependence and the interplay between consumptive and non‐consumptive effects of predators

1. Density‐ and trait‐mediated indirect interactions (DMIIs and TMIIs, respectively) in food chains play crucial roles in community structure and processes. However, factors affecting the relative strength of these interactions are poorly understood, including in widespread and important freshwater rice ecosystems. 2. We studied the strength of DMIIs and TMIIs in a food chain involving a predator (the Reeve’s turtle Chinemys reevesii), its herbivorous prey (the apple snail Pomacea canaliculata) and a plant (rice Oryza sativa) in outdoor containers simulating rice fields. We also evaluated consumptive and non‐consumptive effects of the predator on the snail. We removed a fixed proportion of snails every 2 days to simulate prey consumption and introduced a caged turtle that was fed daily with snails to simulate non‐consumptive effects. 3. Direct consumptive effects increased growth of the remaining snails and their per capita feeding rate. Moreover, consumptive and non‐consumptive effects, and their interaction, affected the proportion of snails buried in the soil. This interaction was presumably because increasing food availability per snail induced their self‐burying behaviour. 4. Both DMIIs and TMIIs affected the number of rice plants remaining, whereas their interaction term was not significant. 5. In summary, density dependence and interactions between consumptive and non‐consumptive effects influenced snail growth and behaviour, respectively. However, no cascading effects of these complicated interactions on rice plants were detected.     

Responses of four submerged macrophytes to freshwater snail density (Radix swinhoei) under clear‐water conditions: A mesocosm study

Macrophytes play a key role in stabilizing clear‐water conditions in shallow freshwater ecosystems. Their populations are maintained by a balance between plant grazing and plant growth. As a freshwater snail commonly found in shallow lakes, Radix swinhoei can affect the growth of submerged macrophytes by removing epiphyton from the surface of aquatic plants and by grazing directly on macrophyte organs. Thus, we conducted a long‐term (11‐month) experiment to explore the effects of snail density on macrophytes with distinctive structures in an outdoor clear‐water mesocosm system (with relatively low total nitrogen (TN, 0.66 ± 0.27 mg/L) and total phosphorus (TP, 36 ± 20 μg/L) and a phytoplankton chlorophyll a (Chla) range of 14.8 ± 4.9 μg/L) based on two different snail densities (low and high) and four macrophyte species treatments (Myriophyllum spicatum, Potamogeton wrightii, P. crispus, and P. oxyphyllus). In the high‐density treatment, snail biomass and abundance (36.5 ± 16.5 g/m² and 169 ± 92 ind/m², respectively) were approximately twice that observed in the low‐density treatment, resulting in lower total and aboveground biomass and ramet number in the macrophytes. In addition, plant height and plant volume inhabited (PVI) showed species‐specific responses to snail densities, that is, the height of P. oxyphyllus and PVI of M. spicatum were both higher under low‐density treatment. Thus, compared with low‐density treatment, the inhibitory effects of long‐term high snail density on macrophytes by direct feeding may be greater than the positive effects resulting from epiphyton clearance when under clear‐water conditions with low epiphyton biomass. Thus, under clear‐water conditions, the growth and community composition of submerged macrophytes could be potentially modified by the manual addition of invertebrates (i.e., snails) to lakes if the inhibitory effects from predatory fish are minor.     

Feeding and growth of native,invasive and non-invasive alien apple snails (Ampullariidae) in the United States: Invasives eat more and grow more

The United States hosts one native and five non-native species of aquatic apple snails (Ampullariidae). All are currently found in or around the Everglades in Florida. Two of these introduced species have devastated wetlands in Southeast Asia, but little is known about how they may impact the Everglades. To evaluate potential impacts of introduced apple snails relative to the native species, we investigated plant species preference, consumption rates, growth rates, and growth efficiencies in five introduced and the single native species across eight native macrophytes common in the Everglades. Three of the non-native snails are invasive, one has shown no tendency to expand, and one appears to have minimal direct impact on macrophytes due to its diet. All snails exhibited similar feeding preferences, with Utricularia sp. being the most preferred, Bacopa caroliniana, Sagittaria latifolia, and Nymphaea odorata being of intermediate preference, and Eleocharis cellulosa, Pontederia cordata, Panicum hemitomon and Typha sp. being least preferred (avoided as foods). Consumption and growth was minimal for P. diffusa on all macrophytes. On Utricularia sp. and Bacopa caroliniana, the invasive species Pomacea insularum and P. canaliculata tended to eat more, grow more, and have higher conversion efficiencies than the native P. paludosa or the non-invasive P. haustrum. These contrasts were more often significant for P. insularum than for P. canaliculata. The greater rates of expansion by the invasive species may derive from their enhanced feeding and growth rates.     

Induced chemical defenses in a freshwater macrophyte suppress herbivore fitness and the growth of associated microbes

The freshwater macrophyte Cabomba caroliniana induces a chemical defense when attacked by either the crayfish Procambrus clarkii or the snail Pomacea canaliculata. Induction by either consumer lowers the palatability of the plant to both consumers. When offered food ad libitum, snails feeding on non-induced C. caroliniana grew 2.6–2.7 times more than those feeding on induced C. caroliniana. Because snails fed less on induced plants, this could be a behavioral effect (reduced feeding), a physiological effect of the induced metabolites on the consumer, or both. To assess these possibilities, we made artificial diets with lipid extracts of induced versus non-induced C. caroliniana and restricted control snails to consuming only as much as treatment snails consumed. Growth measured as shell diameter was significantly lower on the diet containing extract from induced, as opposed to non-induced, plants; change in snail mass was more variable and showed a similar, but non-significant, trend. Thus, snails may reduce feeding on induced plants to avoid suppression of fitness. The induced defenses also suppressed growth of co-occurring microbes that might attack the plant through herbivore-generated feeding scars. When two bacteria and three fungi isolated from C. caroliniana surfaces were cultured with the lipid extract from induced and non-induced C. caroliniana, both extracts inhibited the microbes, but the induced extract was more potent against three of the five potential pathogens. Thus, induced plant defenses can act against both direct consumers and microbes that might invade the plant indirectly through herbivore-generated wounds.     

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.     

Zebra mussels decrease burrowing ability and growth of a native snail, <Emphasis Type="Italic">Campeloma decisum</Emphasis>

Invasive species can drive native organisms to extinction by limiting movement and accessibility to essential resources. The purpose of this study was to determine if zebra mussels (Dreissena polymorpha) affect the burrowing ability and growth rate of a native snail, Campeloma decisum. Snails with and without zebra mussels were collected from Douglas Lake, MI, and burrowing depths were studied in both the laboratory and Douglas Lake. Growth rates were calculated as the amount of shell growth from 2004 to 2005. Both the tendency of snails to burrow and the depth to which they burrowed into the substrate were significantly decreased by the presence of zebra mussels on snail shells in both laboratory and lake experiments. There was no difference in the percentage of snails that exhibited growth as a function of zebra mussel presence. However, for those snails that grew, there was a 50% higher growth rate for snails without zebra mussels compared to snails with zebra mussels. These negative effects of zebra mussels on growth and burrowing ability will likely lead to decreases in snail population densities in the future. Handling editor: S. Wellekens     

The effect of calcium and pH on Florida apple snail, Pomacea paludosa (Gastropoda: Ampullariidae), shell growth and crush weight

Pomacea (Ampullariidae) snails, commonly referred to as apple snails, serve as prey for many freshwater-dependent predators, and some species are highly invasive. Identifying limits to apple snail distribution and abundance are pertinent to understanding their ecology. Calcium (Ca²⁺) availability and pH generally influences freshwater snail populations, yet scant data exist for Pomacea snails. We measured 6-week change in shell length (ΔSL) in P. paludosa in two laboratory experiments with varying Ca²⁺ and pH levels. ΔSL was significantly higher in ≥28 mg Ca²⁺/l compared with treatments ≤14 mg/l. Snails from populations living in low Ca²⁺/pH waters did not appear genetically predisposed at growing faster in these conditions. Smallest ΔSL was in snails treated with 3.6 mg Ca²⁺/l and pH < 6.5 water; these snails had signs of shell erosion. Shell crush weights (CWs) were lowest for snails grown in the lowest Ca²⁺/pH treatment. Smaller shells and lower CWs have implications for predation vulnerability and reproductive success. Our results are consistent with reports associating relatively low snail densities with relatively low Ca²⁺/pH waters, and they are consistent with the geographic distribution of P. paludosa as related to the underlying water chemistry as influenced by geology.     

Host-specific performance and host use in the kleptoparasitic marine snail<Emphasis Type="Italic"> Trichotropis cancellata</Emphasis>

Generalist parasites may disproportionately use certain hosts because of different benefits associated with each host species. I measured the growth rate of the marine snail Trichotropis cancellata, a facultative kleptoparasite that can suspension feed and steal food, on different hosts to determine the relative nutritional benefits of each host. The variation in tentacle (feeding structure) area among the hosts studied had the potential to provide parasitic snails with different amounts of nutrition for growth. In field experiments, suspension-feeding snails isolated from potential hosts grew at a similar rate to snails on brachiopods and significantly more slowly than snails on the following polychaete worms: Serpula columbiana (Serpulidae), Pseudopotamilla ocellata (Sabellidae), Schizobranchia insignis (Sabellidae), and Eudistylia vancouveri (Sabellidae). However, choice among worm hosts affected snail growth rates only in the fall, when phytoplankton levels are low. At this time, snails parasitizing the sabellids Schizobranchia and Eudistylia grew more quickly than snails on Serpula. In the spring and summer, with high levels of phytoplankton, Trichotropis grew at similar rates on all worm species tested. Trichotropis spent approximately the same time stealing food from each worm host species, >50% of the time the worms had their tentacles extended (the difference among hosts was not significant). This finding demonstrates that the similarity of snail growth rates on different worm species is not due to the snails compensating for poor hosts (worms that provide food at a slower rate) by spending more time stealing food. Snails in choice experiments preferred live Serpula to empty Serpula tubes, indicating that at least some of the cue(s) snails use to identify hosts are derived from living host tissues. In choice racks containing live Serpula and live Schizobranchia, snails did not choose one host worm significantly more often than the other. Because Trichotropis grows faster on sabellids than serpulids in the fall, I predicted that snails in nature would infect sabellids more often than other species. However, snails were usually distributed randomly among host species. In the few cases where the snails showed a significant preference among host species, proportionally more snails were found on serpulids than on sabellids or sabellarids. This study is the first to quantify under natural conditions the growth benefits of a kleptoparasite across the range of possible hosts, and implies that factors other than growth rate influence host choice specificity in the marine kleptoparasite T. cancellata.     

Life history traits determine the differential vulnerability of native and invasive apple snails (<Emphasis Type="Italic">Pomacea</Emphasis> spp.) to a shared juvenile-stage predator

The vulnerability of gastropods to their predators varies with life history traits such as morphology, body size, behavior, and growth rates as well as predator size. A recent study suggested that the invasive apple snail, Pomacea maculata, was considerably more vulnerable to crayfish predators than the native Florida apple snail, P. paludosa. The difference was hypothesized to be caused by the relatively small hatchling size of P. maculata. To test this hypothesis, we conducted a series of feeding assays designed to quantify maximum feeding rates and selective foraging of crayfish on apple snails. The rate at which crayfish killed individual P. maculata (i.e., kill rates) decreased with snail size, and kill rates on both species increased with crayfish size. Kill rates on juvenile P. maculata were higher than kill rates on size-matched hatchling P. paludosa, and crayfish fed selectively on P. maculata when offered mixed groups of size-matched snails. Further analyses revealed that hatchling P. paludosa possess shells 1.8× heavier than size-matched P. maculata suggesting differences in vulnerability to crayfish were consistent with interspecific differences in shell defenses. Differences in hatchling size and defensive traits in combination make crayfish kill rates on hatchling P. maculata approximately 15.4× faster than on hatchling P. paludosa, but the relative contribution of hatchling size to differences in apple snail vulnerability was >3× greater than the contribution of defensive traits.     

Factors affecting the distribution and abundance of the commensal <Emphasis Type="Italic">Temnocephala iheringi</Emphasis> (Platyhelminthes: Temnocephalidae) among the southernmost populations of the apple snail <Emphasis Type="Italic">Pomacea canaliculata </Emphasis>(Mollusca: Ampullariidae)

Temnocephala iheringi is the most common temnocephalan inhabiting the mantle cavity of the apple snail Pomacea canaliculata, a freshwater neotropical gastropod that has become a serious rice pest in Southeastern Asia. T. iheringi has been recorded from Mato Grosso (Brazil) to water bodies associated with the Río de la Plata river (Argentina). During an extensive survey in the southern limit of the native area of P.␣canaliculata the presence of T. iheringi eggs was recorded in several apple snail populations, extending the known distribution of the commensal more than 400 km southwards. The aim of this study was to understand the factors affecting the distribution and abundance of T. iheringi among populations of P.␣canaliculata. Only 23% of the apple snail populations inhabiting streams harboured temnocephalans while the occurrence among lentic ones was 71%. T. iheringi was found mostly in populations of apple snails living in non-alkaline sites and where snails attaining sizes larger than 4 cm were very common. The prevalence of the temnocephalans in lentic populations was higher than 90%. The number of eggs on the shell (not including the umbilicus) ranged between 0 and 470 and was different among populations of P.␣canaliculata. The prevalence and number of eggs were lower in the lotic populations, except for a stream population immediately downstream of a lake with commensals. There was no difference between males and females of P. canaliculata neither in the prevalence nor in the number of eggs on the shell. The southernmost population of the world of P. canaliculata harbours commensals that tolerate cold winter water temperatures (4–5 °C) as well as its host. On the other hand, T. iheringi was found only in sites with bicarbonate concentrations lower than 6.6 meq l⁻¹, suggesting that the tolerance of the commensal is very much lower than that of the apple snail (up to 9.95 meq l⁻¹). The number of worms inside each snail or the life history variation of P. canaliculata could explain the influence of the size of the snails on the occurrence of T. iheringi. In the big-sized snails, where the number of commensals is higher, the probability of survival of at least one worm is also higher, specially during the hibernation period, when crawling and feeding are null and snails remain buried. On the other hand, P. canaliculata snails from lentic populations are generally bigger and mostly iteroparous, while those inhabiting streams are smaller and semelparous. In these populations the snails have access to mate only with snails of their same cohort, while in iteroparous populations they can copulate with individuals of other cohorts, allowing the inter-generation transmission of worms and the long term persistence of the population of commensals.     

Wetland Restoration and Invasive Species: Apple snail (Pomacea insularum) Feeding on Native and Invasive Aquatic Plants

The apple snail Pomacea insularum is an aquatic invasive gastropod native to South America that has the potential to cause harm to aquatic ecosystems, wetland restoration, and agriculture. To predict the potential impact of this snail on aquatic ecosystems, we tested the feeding rate of P. insularum , under laboratory nonchoice experiments, for 3 species of invasive macrophytes and 13 species of native aquatic plants that are important for wetland restoration and health. High levels of consumption were recorded for four native species ( Ceratophyllum demersum , Hymenocallis liriosme , Ruppia maritima , and Sagittaria lancifolia ) and three invasive species ( Colocasia esculenta , Alternanthera philoxeroides , and Eichhornia crassipes ). In contrast, less than 10% of the biomass of Spartina alterniflora , Scirpus californicus , Thalia dealbata , and Typha latifolia was consumed by P. insularum over the test period. The palatability of macrophytes was negatively correlated with dry matter content, making our results generalizable to all regions where this invader may be present. Based on our results, wetland restoration in areas invaded by P. insularum should focus on emergent structural species with low palatability. Apple snails should not be considered as agents of biocontrol for invasive plants; although apple snails fed on invasive plants at a high rate, their consumption of many native species was even greater.