Introduction,distribution, spread,and impacts of exotic freshwater gastropods in Texas

We examined the patterns of distribution, vectors of introduction, and potential ecological impacts of freshwater exotic species in Texas over the last 45 years. Currently, five species of exotic gastropods are established: channeled-type applesnail (Pomacea insularum), red-rim melania (Melanoides tuberculatus), quilted melania (Tarebia granifera), giant rams-horn snail (Marisa cornuarietis), and Chinese mysterysnail (Cipangopaludina chinensis). In contrast to the northern part of the US, where shipping appears to be the most important vector for the introduction of aquatic invasive species, aquarium and ornamental trade dominated among unintentional vectors of introduction of all freshwater exotics in Texas, resulting in different patterns of distribution, spread, and ecological impacts. The rate of spread of exotic gastropods in Texas varied from 39 waterbodies colonized over 18 years for P. insularum to only three waterbodies during last 45 years for C. chinensis. Four of five exotic gastropods were found in highly vulnerable aquifer-fed springs and rivers, which contain numerous endemic and endangered species. The fifth species, Pomacea insularum, is an agricultural pest. Potential negative ecological effects of exotic gastropods include impacts on wetlands and wetland restoration, competitive exclusion of native snails, and the introduction of exotic parasites, trematodes, which could infect fish and waterfowl, including federally protected species. Aquifer springs with stable temperature regimes are refuges for both cold and warm intolerant species. Handling editor: D. Dudgeon     

The abundance of an invasive freshwater snail Tarebia granifera (Lamarck, 1822) in the Nseleni River,South Africa

The invasive freshwater snail Tarebia granifera (Lamarck, 1822) was first reported in South Africa in 1999 and it has become widespread across the country, with some evidence to suggest that it reduces benthic macroinvertebrate biodiversity. The current study aimed to identify the primary abiotic drivers behind abundance patterns of T. granifera, by comparing the current abundance of the snail in three different regions, and at three depths, of the highly modified Nseleni River in KwaZulu-Natal, South Africa. Tarebia granifera was well established throughout the Nseleni River system, with an overall preference for shallow waters and seasonal temporal patterns of abundance. Although it is uncertain what the ecological impacts of the snail in this system are, its high abundances suggest that it should be controlled where possible and prevented from invading other systems in the region.     

Benthic assemblages of wetlands invaded by Tarebia granifera (Lamarck, 1822) (Caenogastropoda: Thiaridae) in the iSimangaliso Wetland Park,South Africa

Tarebia granifera is a freshwater/estuarine gastropod invading many tropical and sub-tropical areas around the world. This snail is native to southeast Asia and was accidentally introduced into South Africa during the last decade. The current study investigated shallow-water benthic assemblages of different invaded and uninvaded localities across locations spanning a large range of environmental conditions in the iSimangaliso Wetland Park. Using a correlation-based approach, we found that native benthic assemblages were more closely associated with environmental conditions than with densities of T. granifera. However, there were significant negative correlations between T. granifera abundance and Shannon Diversity at two of the invaded locations. This alien species has successfully invaded, and become dominant in, different types of water bodies with different assemblage compositions and physico-chemical characteristics, ranging from freshwater ponds to saline estuaries and lakes. The current data set is presented as an essential baseline for future studies. It is recommended that future work focuses on specific localities, in order to determine if changes in diversity are driven by non-native species or by other disturbances (e.g., climate change).     

Neritid and thiarid gastropods from French Polynesian streams: how reproduction (sexual, parthenogenetic) and dispersal (active, passive) affect population structure

• 1 The streams of French Polynesia contain several species of Neritidae and Thiaridae (Mollusca: Gastropoda). The neritids are dioecious and amphidromous with a freshwater adult stage and a poorly known, marine larval stage. The thiarids are parthenogenetic and viviparous, and rely on passive dispersal for colonisation of new habitats.
• 2 Populations of the neritid Clithon spinosus and the thiarids Melanoides tuberculata and Thiara granifera were analysed using mitochondrial DNA sequences from COI to compare the population structure of the snails at three different scales: between streams (N = 9), between islands (N = 4), and between age and distance of paired islands.
• 3 The amphidromous C. spinosus showed no evidence of genetic isolation at any of the scales tested (F_st values 0.02). Parsimony analyses resulted in two haplotype clusters separated by a three‐step segment, which were not linked to geographic isolation. The larval phase of C. spinosus is most likely a long‐lived planktotroph and a very effective disperser.
• 4 Two haplotypes of M. tuberculata, separated by 16 base pairs, were found. Both haplotypes were found in snails on all islands, and individuals representing both were often collected in the same habitat. One haplotype of T. granifera was found. M. tuberculata has the characteristics of the ‘general‐purpose genotype’ of clonal population structure and although it relies on passive dispersal, it has colonised nearly all freshwater habitats on the islands.

     

Experimental test of the Invasional Meltdown Hypothesis: an exotic herbivore facilitates an exotic plant,but the plant does not reciprocally facilitate the herbivore

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Stable isotope evidence for dietary overlap between alien and native gastropods in coastal lakes of northern KwaZulu-Natal, South Africa

Background

Tarebia granifera (Lamarck, 1822) is originally from South-East Asia, but has been introduced and become invasive in many tropical and subtropical parts of the world. In South Africa, T. granifera is rapidly invading an increasing number of coastal lakes and estuaries, often reaching very high population densities and dominating shallow water benthic invertebrate assemblages. An assessment of the feeding dynamics of T. granifera has raised questions about potential ecological impacts, specifically in terms of its dietary overlap with native gastropods.

Methodology/Principal Findings

A stable isotope mixing model was used together with gut content analysis to estimate the diet of T. granifera and native gastropod populations in three different coastal lakes. Population density, available biomass of food and salinity were measured along transects placed over T. granifera patches. An index of isotopic (stable isotopes) dietary overlap (IDO, %) aided in interpreting interactions between gastropods. The diet of T. granifera was variable, including contributions from microphytobenthos, filamentous algae (Cladophora sp.), detritus and sedimentary organic matter. IDO was significant (>60%) between T. granifera and each of the following gastropods: Haminoea natalensis (Krauss, 1848), Bulinus natalensis (Küster, 1841) and Melanoides tuberculata (Müller, 1774). However, food did not appear to be limiting. Salinity influenced gastropod spatial overlap. Tarebia granifera may only displace native gastropods, such as Assiminea cf. ovata (Krauss, 1848), under salinity conditions below 20. Ecosystem-level impacts are also discussed.

Conclusion/Significance

The generalist diet of T. granifera may certainly contribute to its successful establishment. However, although competition for resources may take place under certain salinity conditions and if food is limiting, there appear to be other mechanisms at work, through which T. granifera displaces native gastropods. Complementary stable isotope and gut content analysis can provide helpful ecological insights, contributing to monitoring efforts and guiding further invasive species research.     

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     

Comparisons of isotopic niche widths of some invasive and indigenous fauna in a South African river

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Coexistence in the intertidal: interactions between the non‐indigenous New Zealand mud snail Potamopyrgus antipodarum and the native estuarine isopod Gnorimosphaeroma insulare

Nonindigenous species (NIS) that achieve high densities in their recipient communities are expected to have negative effects on native competitors. However, recent reviews have shown that competitors, unlike predators and parasites, have rarely been documented to cause extinctions. There is a need for better understanding of competitive interactions between nonindigenous species and native competitors across systems where NIS reach high densities. In this paper we examine competitive interactions between the exotic, invasive New Zealand mud snail Potamopyrgus antipodarum and native estuarine invertebrates. P. antipodarum is often described as a freshwater snail but can occur in brackish water as well; it has been established in the Columbia River Estuary for over ten years. We addressed competition in this estuarine system using three approaches: field surveys of the benthic invertebrate community, stable isotope analysis of overlap in resource use by common invertebrates, and a laboratory competition experiment that tested the strength of competition between P. antipodarum and the isopod, Gnorimosphaeroma insulare, which overlap in resource use. We found no evidence of negative competitive impacts of P. antipodarum on native benthic invertebrates in this estuarine system. Densities of P. antipodarum and common native epibenthic invertebrates are positively correlated in the intertidal, likely due to shared habitat preferences. The competition experiment showed that the effect of interspecific competition from P. antipodarum on the foraging and survival of G. insulare was significantly less than the effect of intraspecific competition between isopods. The presence of the isopod G. insulare reduced foraging in P. antipodarum, but this change did not result in reduced survival and growth of snails. Hence, interspecific competition is weak despite high densities achieved by the invading species. Finally, we discuss several possible explanations for the observation that P. antipodarum does not have an obvious negative competitive impact on native benthic invertebrates.     

An experimental investigation of interactions in snail-macrophyte-epiphyte systems

Summary An experimental investigation under field conditions of enclosures containing freshwater pulmonate snails, the macrophyteCeratophyllum demersum and epiphytes, produced evidence of beneficial interactions.Ceratophyllum growth, measured in terms of stem length, numbers of leaf-nodes and growing tips and leaf survival was significantly enhanced in the presence of snails. This effect was attributed to the increased availability of plant nutrients of snail origin, such as phosphates and ammonia, as well as to the snails' action as “cleaning symbionts” in reducing the density of bacterial and algal epiphyton potentially deleterious to macrophytes. Principal component analysis revealed both seasonal and treatment effects of snail grazing on algal epiphyton. Small adnate algal species (e.g.Cocconeis placentula) survived grazing and benefited from the removal of larger, competitor, species. Snail densities increased in all treatments, despite high (86%) juvenile mortality. It is concluded that freshwater pulmonate snails are strong interactors in lentic habitats, enhancing the growth ofCeratophyllum and producing characterisic epiphyte communities. This benefits not only the snails, but also the plants and epiphytes that are associated with them. Thus the interactions between these component parts of the community can be considered as mutualistic.     

Distribution,movement, and microhabitat use of the introduced predatory snail Euglandina rosea in Hawaii: implications for management

The purposeful introduction of the land snail Euglandina rosea, which feeds exclusively on snails and slugs, has been implicated as a major factor in the decline of diverse Pacific island land snail faunas. We report on the distribution, movement patterns, and microhabitat preferences of E. rosea in a gulch in the Waianae Mountains, Oahu, Hawaii, because such data will help focus management actions at a local scale to protect native snail populations in areas where E. rosea is established. The Waianae Mountains harbor many endangered or threatened snails, most currently found in isolated habitat patches near the ridges. Conversely, most living individuals (28/29) and shells (46/56) of E. rosea were collected within the gulch, which supported higher densities of other native and non‐native snails, and was cooler and more moist than the ridges. Thirteen individuals of E. rosea were tracked (eight directly using a bobbin and thread method, and five indirectly by mark–recapture); most (10/13) moved on average <2.5 m per week (range 0.1–25.21 m), and all stayed within the gulch. Members of E. rosea preferred leaf litter over open, fern/shrub, or wood microhabitats. There were large differences in the population density of E. rosea over small spatial scales, indicating that there may be places where native snail populations could persist even in areas where populations of E. rosea are established. Identifying areas with differing population densities of E. rosea is critical for not only understanding why some native snail species may be more vulnerable to extinction, but also to locate areas where predation pressure is low and conservation efforts will be most likely to succeed.     

Growth in a freshwater snail under laboratory conditions in relation to eutrophication

Shell growth in the snail Potamoryrgus jenkinsi (Prosobranchia, Hydrobiidae) was measured under laboratory conditions. Individuals of populations from two lakes differing in trophic status were kept in water with sandy substrates from each environment. Growth was faster in the environment of the more eutrophic lake, regardless of the origin of the snails; this held in particular for the smallest size class.     

Population structure of Placostylus hongii (Gastropoda: Paryphantidae) on the Poor Knights Islands

Abstract

Populations of the native land snail Placostylus hongii (Lesson, 1830) on the Poor Knights Islands were investigated during visits in 1976, 1977, and 1978. Marking and recapture of individual snails showed that linear growth of the shell ceased when a lip was formed around the outer margin of the aperture. Measurements of snails suggested that increase in outer lip width was a good indication of age, but that the rate of lip thickening varied between populations. Large Placostylus showed a restricted range of movement. Both tagging studies and random sampling revealed that mortality was size specific. Most small snails (<60 mm shell length) located were dead, whereas most larger snails were alive. Population densities of living snails averaged 2.2 individuals per 0.25 m² in the study area. The local distribution was patchy owing to the tendency of snails to be associated with specific features of the vegetation.     

Variation of snail's abundance in two water bodies harboring strains of Pseudosuccinea columella resistant and susceptible to Fasciola hepatica miracidial infection, in Pinar del Río Province, Cuba

The abundance of freshwater snails in two rural sites of Pinar del Río, Cuba, which harbor Pseudosuccinea columella susceptible and resistant to miracidia of Fasciola hepatica was followed for one year. Susceptible snails were found in the most anthropic site (IPA) whereas the resistant population inhabited the most preserved one (El Azufre). Only two snail species coexisted with P. columella at IPA site (Physa cubensis and Tarebia granifera) while five species were found at El Azufre, including an endemic from that province (Hemisinus cubanianus). Populations of both resistant and susceptible snails showed stable densities throughout the year, although the susceptible strain attained higher abundance. The highest densities were observed in April-May 2004 for the susceptible population whereas the resistant strain attained its highest abundance in January 2004. No record of Fossaria cubensis was made and the thiarid T. granifera occurred only at low densities. One of the sampled sites (IPA) meets all the conditions for the first report of P. columella naturally infected with larvae of F. hepatica.     

Density-dependent growth and reproduction of the apple snail, Pomacea canaliculata: a density manipulation experiment in a paddy field

To examine density dependence in the survival, growth, and reproduction of Pomacea canaliculata, we conducted an experiment in which snail densities were manipulated in a paddy field. We released paint-marked snails of 15–20 mm shell height into 12 enclosures (pens) of 16 m² at one of five densities – 8, 16, 32, 64, or 128 snails per pen. The survival rate of released snails was 95% and was independent of snail density. The snail density had a significant effect on the growth and egg production of individual snails. This density dependence may have been caused by reduced food availability. The females at high density deposited fewer and smaller egg masses than those at low density, and consequently produced fewer eggs. The females at densities 8 and 16 deposited more than 3000 eggs per female, while the females at density 128 oviposited only 414 eggs. The total egg production per pen was, however, higher at higher snail density. The survival rates of juvenile snails were 21%–37% and were independent of adult density. The juvenile density was positively correlated with the total egg production per pen and hence was higher at higher adult density. However, the density of juveniles larger than 5 mm in shell height, i.e., juveniles that can survive an overwintering period, was not significantly different among density treatments. These results suggest that snail density after the overwintering period is independent of the density in the previous year. Thus, density dependence in growth and reproduction might regulate the population of P. canaliculata in paddies. Received: October 23, 1998 / Accepted: July 16, 1999     

Detrital Fruit Processing in a Hawaiian Stream Ecosystem1

In temperate forested streams, fruit from riparian trees is generally a minor and seasonal component of the allo‐chthonous detritus. In contrast, riparian fruit input to tropical streams is often high and continuous. Detrital fruit is abundant in some forested Hawaiian streams compared to other forms of riparian detritus, and rates of leaf litter processing by macroscopic invertebrates are very low. These observations suggested that fruit is an important food resource for detritivores. A microcosm system was used to measure the rates at which two common detrital fruits, guava and mango, were processed by two common detritivores, the prawn Macrobrachium lar and the gastropod Tarebia granifera. Comparisons of fruit weight loss rates normalized by detritivore weight indicated that M, lar processed guavas at significantly higher rates than T. granifera, differences in rates of mango processing by M. lar and T. granifera were not significant. Microcosms containing both M. lar and T. granifera were used to test for interactions between the invertebrates that affected rates of mango processing. No interspecific interactions were detected. A field study was conducted in Kaiwiki Stream, Island of Hawaii, to determine rates of detrital fruit input and export. Detrital fruit was supplied to the study area year‐round, with peaks corresponding to summer and autumn fruiting seasons. Guavas and mangos accounted for 85 percent of the fruit biomass entering the stream and 92 percent of the fruit exported from the stream. Mean daily export rates of guava were 7 percent of input, and export rates of mango were 5 percent of input. These measurements suggested that most of the fruit entering the stream is retained and comprises a substantial food resource for detritivores. Comparisons of the biomass‐specific rates at which M. lar and T. granifera processed mangos and guavas with the rates at which mangos and guavas entered Kaiwiki Stream suggested that these invertebrates can process most of the detrital fruit in the stream.     

Neighborhood‐contingent indirect interactions between native and exotic plants: multiple shared pollinators mediate reproductive success during invasions

Native and exotic plants can influence one another's fecundity through their influence on shared pollinators. Specifically, invasion may alter abundance and composition of local floral resources, affecting pollinator visitation and ultimately causing seedset of natives in more‐invaded and less‐invaded floral neighborhoods to differ. Such pollinator‐mediated effects of exotic plants on natives are common, but native and exotic plants often share multiple pollinators, which may differ in their responses to altered floral neighborhoods. We quantified pollinator‐mediated interactions between three common forbs of western Washington prairies (native Microseris laciniata and Eriophyllum lanatum and European Hypochaeris radicata) in three floral neighborhoods: 1) high native and low exotic floral density, 2) high exotic floral density and low native density, and 3) experimentally manipulated low exotic floral density. Pollinator visitation rates varied by floral neighborhood, plant species identity, and their interaction for all three plant species. Similarly, pollinator functional groups (eusocial bees, solitary bees, and syrphid flies) contributed differing proportions of total visitation to each species depending upon neighborhood context. Consequently, in exotic neighborhoods H. radicata competed with native M. laciniata, reducing seed set, while simultaneously facilitating visitation and seed set for native E. lanatum. Seed set of H. radicata was also highest in exotic neighborhoods (with high densities of conspecifics), raising the possibility of a positive feedback between exotic abundance and success. Our results suggest that the outcome of indirect interactions between native and exotic plants depends on the density and the composition of the floral neighborhood and of the pollinator fauna, and on context‐dependent pollinator foraging.     

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.     

Competition and facilitation within and between a snail and a mayfly larva and the effect on the grazing process

We studied the competitive effects within and between two taxonomically distant freshwater herbivores, a snail and a mayfly, common in Swedish lakes, Lymnaea peregra and Cloeon dipterum, respectively, and their effect on grazing in a laboratory experiment. The experimental set-up consisted of 2-l aquaria, each containing a periphyton covered tile. Intra- and interspecific effects were tested by increasing the density of one species at a time in four different treatments, (1) snails (intraspecific treatment), (2) mayflies (intraspecific treatment), (3) mixed-snails (interspecific treatments, snails kept constant) and (4) mixed-mayflies (interspecific treatments, mayflies kept constant). Intraspecific competition affected both snails and mayflies negatively, i.e. increasing mortality with increasing con-specific density. Furthermore, there was a decrease in snail growth with increasing snail density. In the mixed-species treatments both species changed their microhabitat use indicating interspecific competition. Despite this, we also found a positive effect of mayfly density on snail growth, most likely due to indirect commensalism. No density-dependent effect of grazing on periphyton was found, probably due to interference competition between grazers. However, there was a significant difference in periphyton biomass, due to species composition of grazers. Irrespective of their densities, if they co-existed, the two grazer species decreased the periphyton biomass significantly compared with both single-species treatments. We considered this as a joint action of facilitation and interaction. Our results suggest that competition can be an important structuring factor in macroinvertebrate communities and that species composition can be significant for ecosystem processes within lentic environments.     

Snail populations in arctic lakes: competition mediated by predation?

Summary For 2 species of snails in arctic Alaskan lakes, I studied the patterns of snail distribution with respect to habitat, distribution of predatory fish, and the potential for interspecific competition. The snails Lymnaea elodes and Valvata lewisi co-exist in these arctic lakes, either in the presence of lake trout, Salvelinus namaycush, or in the absence of predation. Intensive sediment core sampling of Toolik Lake and Lake N-2, with trout and lacking trout, respectively, showed that the smaller snail, Valvata, was abundant in Toolik but ocurred at very low densities in Lake N-2. On the open sediments of lakes containing trout, diver surveys revealed very low densities of adult Lymnaea (0.12±0.12/m²), but similar surveys in lakes without trout revealed much higher densities of adult Lymnaea (7.1±1.8/m²). A survey of 14 lakes indicated that adult Lymnaea grew to a smaller mean size in lakes with trout than in lakes which lacked trout.In laboratory and field experiments, the presence of Lymnaea lowered the fecundity of Valvata. Laboratory experiments also showed that Lymnaea fecundity was enhanced by the presence of Valvata. Enhancement was not due to predation by Lymnaea on Valvata eggs or young. The observed patterns of distribution and abundance in the absence of trout, combined with results from laboratory experiments, are consistent with the hypothesis that competitive and facilitative interactions control the population dynamics of the two snails. The distribution and abundance patterns of snails where trout are present suggest that trout predation rather than competition controls snail population dynamics in lakes containing trout.