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     

The effects of predation by the mummichog, Fundulus heteroclitus (L.), on the abundance and distribution of the salt marsh snail, Melampus bidentatus (Say)

Enclosure and exclosure experiments were conducted in Canary Creek marsh to examine how predation by a killifish, Fundulus heteroclitus (L.), affects the abundance and size distribution of the salt marsh snail, Melampus bidentatus (Say). Enclosures (7.6 × 19.7 m) were stocked with Fundulus heteroclitus at densities of one-half normal, normal, twice normal, and four times normal density. Fish exclusion pens were also built. In both years of study, the mean density of snails increased significantly in pens where fish were excluded or their density reduced. During the same period in each year, the density of snails in pens containing higher than normal fish density fell by ≈50%.

Fish density also affected the size distribution of snails within pens. In both years, mean shell length of snails in the pen with the highest density of fish was significantly greater, and mean shell length of snails within fish exclusion pens was significantly lower than in all other treatments. Gape size limitation of F. heteroclitus causes selective predation on small snails and apparently is responsible for the difference in mean shell length among treatments.

Density and size distribution measurements of the natural Melampus bidentatus population in Canary Creek marsh were also taken. Larger snails were found to occur in the low marsh zone, whereas smaller snails occurred in the high marsh zone. Since the high marsh area is flooded less often than the low marsh, and the grass types found in the high marsh zone are known to afford protection from fish predation, the distribution of snails in the marsh is consistent with the idea that fish predation is an important factor influencing the distribution of snails in Canary Creek marsh.     

Effect of immersion at low tide on distribution and movement of the mud snail, Ilyanassa obsoleta (Say), in the upper Bay of Fundy, eastern Canada

Habitat heterogeneity often affects movement behaviours of animals, and consequently their spatial distribution. We evaluated the effect of immersion at low tide on the distribution, fine-scale movement patterns and daily movement patterns of the mud snail Ilyanassa obsoleta on a mudflat in the upper Bay of Fundy, Canada. Mud snails migrate onto intertidal mudflats in the summer, and our field survey showed that their density was higher inside tide pools relative to adjacent areas that are exposed at low tide. Using time-lapse videography, we evaluated the effect of snail size, snail density, and immersion at low tide on fine-scale movement patterns of I. obsoleta. Time until snails stopped moving and burrowed was unaffected by snail size, but snails at low and high densities burrowed somewhat faster than those at intermediate densities. Snail size and snail density had no detectable effect on displacement speed or linearity of displacement. Immersion affected snail movement: snails within tide pools delayed burrowing and traveled in more convoluted paths compared to those on exposed mud. Snails increased their turning angles within tide pools, which is probably the mechanism by which aggregations are formed. We also performed a mark-recapture experiment to compare daily movement patterns of snails released inside and outside tide pools. Snails released in tide pools moved shorter distances, but did not orient themselves differently than snails released outside tide pools. Both groups exhibited significant directionality, moving against the mean water current direction over 24 h. In sum, immersion at low tide affected the behaviour and spatial distribution of snails, resulting in snail aggregations within tide pools. These snail aggregations, in turn, may be a major factor influencing spatial dynamics on mudflats, including causing changes in distribution patterns of the burrowing amphipod Corophium volutator, a dominant inhabitant and key species in the food web of mudflats.     

Changes in the selection differential exerted on a marine snail during the ontogeny of a predatory shore crab

Empirical estimates of selection gradients caused by predators are common, yet no one has quantified how these estimates vary with predator ontogeny. We used logistic regression to investigate how selection on gastropod shell thickness changed with predator size. Only small and medium purple shore crabs (Hemigrapsus nudus) exerted a linear selection gradient for increased shell‐thickness within a single population of the intertidal snail (Littorina subrotundata). The shape of the fitness function for shell thickness was confirmed to be linear for small and medium crabs but was humped for large male crabs, suggesting no directional selection. A second experiment using two prey species to amplify shell thickness differences established that the selection differential on adult snails decreased linearly as crab size increased. We observed differences in size distribution and sex ratios among three natural shore crab populations that may cause spatial and temporal variation in predator‐mediated selection on local snail populations.     

Density and environmental effects on shell size in some sand dune snail populations

Populations ol three species of land snail, Helicella itala. Candidula intersecta and Cochlicella acuta are studied on sand dunes on Coll in the Inner Hebrides.
Population density and mean shell size were estimated on 18 sample sites. The sites were ranked for six environmental factors. Environmental factors which are related to the dune vegetational succession account for much of the variation in the densities of C. intersecta and C. acuta . The density of H. itala , however, shows no strong association with these factors. Both snail density and shell size are reactively independent of total soil calcium levels. For each of the three species, mean shell size is negatively associated with population density; the environmental factors account for little of the variation in shell size.
Four possible mechanisms are suggested to account for the variation of shell size with density. It is argued that a direct influence of density on shell size, possibly mediated by mucus conditioning of the environment is the most probable mechanism. There is some evidence to suggest an inter-specific effect whereby C. intersecta density affects H. itala shell size.     

Gastropod Colonization of a Created Coastal Wetland: Potential Influences of Habitat Suitability and Dispersal Ability

We investigated the assumption that populations of epibenthic macroinvertebrates readily establish in created coastal wetlands by quantifying the spatial and temporal patterns of Cerithidea californica (California horn snail) density in a newly created wetland and an adjacent natural area in Mugu Lagoon, southern California, United States, for 3.5 years. The natural and created sites differed in vascular plant cover and sediment grain size, organic content, salinity, and moisture content. Densities of C. californica in the created site changed little during the study period, and were often lower than those in the natural site. The influences of habitat suitability and dispersal limitation on C. californica colonization of the created site varied among snail age classes. Sediment moisture and organic content explained some of the variability in subadult (47%) and adult (55%) density and relative abundance, but none of the variability in juvenile abundance. Adult snail density was also strongly influenced by distance from the natural/created site transition zone. Juvenile and subadult snail densities were not related to distance from the natural site, possibly due to greater dispersal ability. Between‐site differences in C. californica densities and size structure suggested that adult snails were affected by both habitat characteristics and dispersal ability, subadults were influenced to a lesser degree by habitat characteristics, and juveniles were not related to either factor. Accordingly, the influence of habitat characteristics and dispersal ability on created site colonization may change with snail age. Successful restoration of benthic invertebrate communities requires consideration of both habitat characteristics and dispersal ability of the target species, even in created sites in close proximity to natural areas.     

Small- and broad-scale patterns of distribution of the upper-shore littorinid Nodilittorina pyramidalis in New South Wales

Abstract mall-scale (within-shore) and broad-scale (among-shore) patterns of distribution, abundance and size structure of Nodilittorina pyramidalis were measured on a number of shores in New South Wales. This species showed significant patchiness in distribution and size structure. The mean density and mean shell length of snails differed among patches of shore only a few metres apart, among different heights on the shore and from shore to shore. Differences in densities appeared to be strongly correlated to the distribution of particular microhabitats, especially cracks, crevices and pits within the rock surface. This relationship and the presence or absence of these microhabitats appeared to determine the patterns of distribution of this species within and among different shores. The sizes of snails were not correlated with densities of Nodilittorina nor with the coexisting species, Littorina unifasciata, in the populations examined. Processes influencing the distribution and abundance of Nodilittorina were not investigated, but the patterns of distribution found on these shores suggest that recruitment was limited to areas of topographic complexity, ft also seems likely that limited dispersal by the adult snails retain them within restricted patches on the shore.     

Dispersal of the estuarine gastropod Pyrazus ebeninus is only weakly influenced by pneumatophore density

Studies on rocky intertidal gastropods indicate habitat complexity and body size to be major determinants of dispersal patterns. Considerations of effects of habitat complexity and body size on soft sediment gastropods are, however, less common. In neither habitat has the interaction between habitat complexity and body size been considered despite the increasing recognition in the general ecological literature that complexity effects are body-size-dependent. We tested independent and interacting effects of habitat complexity and body size on movement of the mud-whelk, Pyrazus ebeninus, by marking large (61-85 mm) and small (31-55 mm) snails in sites with low and high densities of pneumatophores and determining the distance and direction of their dispersal over periods of 1 week, 2 weeks and 1 month. Contrary to our expectation, we found no effect of pneumatophore density on the distance of snail migration over each of the temporal scales; net distance travelled by snails was determined only by body size and idiosynchratic, site-specific factors. The direction of snail movement was, by contrast, influenced on some temporal scales by both pneumatophore density and snail size. Over 1 week, site effects dominated patterns of movement and neither size of snail nor density of pneumatophore produced statistically significant effects. As the temporal scale increased, effects of size and pneumatophore density became increasingly apparent. Over the 1-month period, large snails at all sites and small snails at sites with high pneumatophore density migrated down the shore, while small snails at sites with low pneumatophore displayed non-directional movement. Thus, overall this study provides only weak support for effects of pneumatophore density on snail movement. In combination with other studies, our results suggest that, in comparison to on rocky shores where habitat complexity has strong effects on the distribution, abundance and behaviour of gastropods in soft-sediment systems habitat complexity is a less important structuring agent.     

Population ecology and fishery of Cittarium pica (Gastropoda: Trochidae) on the Caribbean coast of Costa Rica

The West Indian Topshell Cittarium pica is artisanally collected on rocky shores along the Caribbean Coast of Costa Rica. There are neither data on the state of its exploitation nor exist any regulation of the fishery From October 2000 to March 2001, the population dynamics of this species were studied at an unexploited and two exploited sites to determine the present impact of the fishery on the resource. Average population density with 14 ind./m2 about three times higher at the unexploited than at the exploited sites. Length-frequency histograms showed a strong shift towards smaller specimens at the exploited sites, which is also reflected in significantly higher rates of total mortality (Z = 4.05 and 4.47) when compared to the unexploited site (Z = 1.47). Von Bertalanffy growth parameters were estimated as k = 0.19-0.28 (yr-1) and L infinity = 104 mm. No significant differences were found among sites. From these values a range of the growth performance index phi was computed (phi = 3.31-3.48) which lies at the lower end of the values reported for other tropical marine gastropods. The size at first maturity for both sexes combined was estimated as 29.20 +/- 1.14 mm. Exploitation rates > 0.6 for both exploited sites and a large fraction of small specimens (< 30 mm) in the catches suggest overexploitation and recruitment overfishing. Based on the estimated maximum sustainable yield we recommend regulative measures for the fishery such as a control of a minimum landing size of 40 mm and a closure of the fishery during its reproductive period (from July to November).     

Density and environmental effects on shell size in some sand dune snail populations

Populations ol three species of land snail, Helicella itala. Candidula intersecta and Cochlicella acuta are studied on sand dunes on Coll in the Inner Hebrides. Population density and mean shell size were estimated on 18 sample sites. The sites were ranked for six environmental factors. Environmental factors which are related to the dune vegetational succession account for much of the variation in the densities of C. intersecta and C. acuta. The density of H. itala, however, shows no strong association with these factors. Both snail density and shell size are reactively independent of total soil calcium levels. For each of the three species, mean shell size is negatively associated with population density; the environmental factors account for little of the variation in shell size. Four possible mechanisms are suggested to account for the variation of shell size with density. It is argued that a direct influence of density on shell size, possibly mediated by mucus conditioning of the environment is the most probable mechanism. There is some evidence to suggest an inter-specific effect whereby C. intersecta density affects H. itala shell size.     

Age/size- and time-specific effects of Schistosoma mansoni on energy allocation patterns of its snail host Biomphalaria glabrata

Snail hosts of different ages constitute different resource environments for larval trematodes because each individual has a particular energy budget and energy allocation pattern at the time of infection. Effects of monomiracidial trematode infections on shell growth, body dry weight and reproductive effort of the snail host were compared between controls and infected juvenile and adult snails during the prepatent and patent periods. The results demonstrate phenotypic plasticity in the host response. There is an age/size-specific effect characterized by limitation of growth rate when snails are infected as juveniles and reduction of reproductive effort when snails are infected as adults, and a time-specific effect with early enhancement of growth rate and reproductive effort for infected juvenile and adult snails respectively during prepatency, before reduction and cessation during patency. The parasite is considered as a generalist in its energy exploitation strategy of the snail host, but taking into account the differential host responses observed relative to the host age, the possibility of a host preference for juvenile or adult snails is discussed. Received: 4 February 1997 / Accepted: 14 July 1997     

Variation in adult shell morphology and life-history traits in the land snail Oreohelix cooperi in relation to biotic and abiotic factors

In this study, we explore factors influencing variation in adultshell morphology and life-history characteristics (offspringsize and number) in the ovoviviparous land snail, Oreohelixcooperi, from the Black Hills, South Dakota and Wyoming, USA.To date, no study of shell morphology and life-history characteristicsin ovoviviparous land snails has included a combination of datasetsincluding genetic data, life-history traits, shell morphologyand multiple environmental factors. We report that differencesin shell size among populations are strongly related to meanannual temperature (and the highly correlated variable elevation)and population density (measured as shell density). In addition,shell size varies among populations, despite an apparent lackof population genetic differentiation. Common factors thoughtto influence adult shell size, like precipitation and calciumlevels, do not have a significant effect in this study. Adultsize strongly influences per-clutch reproductive output, withlarger snails having larger and more offspring. As mean annualtemperature and shell density affect adult shell size, theyalso indirectly affect per-clutch output. The results suggestthat a large portion of the life-history variation in O. cooperiis environmentally induced, as has been found in oviparous landsnails and brooding freshwater bivalves. (Received 20 July 2006; accepted 10 January 2007)     

Zur Dynamik exploitierter Populationen vonTisbe holothuriae (Copepoda,Harpacticoida)

In weekly exploited populations of the harpacticoid copepodTisbe holothuriae Humes, effects of exploitation rate on population density, age distribution, growth, as well as numerical and biomass yields were studied under conditions of surplus food supply at 22° C and 30 ‰ S. Increase of exploitation rate lowered the adult plus copepodid density and raised the density of nauplii, thus shifting the age distribution in favour of the nauplii. Total population density was almost equal at exploitation rates of 0.10, 0.50 and 0.70, but lower at rates of 0.30 and 0.90. Experiments, which have not yet been completed, have shown a higher resistance against the toxic effect of cadmium in populations exploited at rates of 0.30 or 0.90; populations exploited at the lower rate displayed a lesser degree of resistance than populations exploited at the rate of 0.90. Variability of densities was least at intermediate exploitation rates, whereas the variability of the age structure and the intrinsic rate of natural increase was less at higher rates, indicating smaller fluctuations in the ratio of the regulating variables mortality and fecundity with increasing exploitation. The assumption is made that an appropriate exploitation rate contributes to the achievement of favourable culture conditions for copepods, which are subjected to intense predation in natural populations. Within the range of the exploitation rates selected, biomass yields of nauplii and adults plus copepodids were approximately proportional to exploitation rate; therefore, the maximum rate sustainable by aTisbe holothuriae population cannot be established. From data on adult numerical yield, however, it may be concluded that exploitation rates exceeding 0.90 will not result in increased productivity.     

The population density effects on the reproductive biology of the snail Bradybaena similaris (Férussac, 1821) (Mollusca, Gastropoda)

The influence of population density on some aspects of the reproductive biology of the snail Bradybaena similaris was studied. Molluscs were maintained under 0.2 (isolated), 0.3, 0.6, 1.0, 1.3 and 1.7 snail/m(2) densities. The animals maintained under 0.3 and 0.6 snail/m(2) showed the lowest numbers of eggs laid/snail, being the highest value observed to the 1.7 snail/m(2). The hatching of the snails maintained under 0.3 snail/m(2) density, begun at the 21st day after laying, and the maximum time required to the hatching was 36 days was observed to the eggs came from snails maintained under the densities 0.6, 1.0, 1.3 snail/m(2), respectively. The highest percentage hatchability (55.56%) was observed to isolated snails. The galactogen content in the albumen gland did not seem to accompany the alterations occurred in the reproduction of B. similaris in response to the different population densities.     

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.     

Microgeographical variation in shell size of the land snail Chondrina clienta

Microgeographical variation in shell morphology of the rock-dwelling land snail Chondrina clienta , collected from 30 sites within an area of 0.5 km² on the island of Öland, Sweden, was examined in relation to its own population density and that of a potential competitor (the land snail Balea perversa ) and to environmental factors. Dispersal of marked individuals averaged 83 cm per year within a stone pile and 291 cm in an area of exposed bedrock. Local population density of active C. clienta ranged from 5 to 794 individuals per m². Shell characters were highly intercorrelated, both within and between populations. Principal component analysis revealed that most of the interpopulational variation could be expressed by the single character of shell height, which ranged from 5.54 to 6.94 mm. In all populations, snails of a given size had the same whorl number. Shell size was not influenced by habitat type (exposed rock surface, stone pile or stone wall) or proportion of calcareous stones within habitat. It was, however, negatively correlated with local population density, indicating intraspecific competition, and positively correlated with the degree of plant cover within the habitat. Analysis of variance revealed additionally a density effect of B. perversa on shell size in C. clienta , probably as a result of interspecific competition. Breeding experiments using C. clienta from different sites and carried out under unifrom conditions caused most of the phenotypic variation to disappear, demonstrating the high phenotypic plasticity of the species.     

不同初始螺类密度对沉水植物刺苦草Vallisneria spinulosa Yan及其附着藻类的影响

大型水生植物及其附着藻类是浅水湖泊中的重要初级生产者。淡水螺类作为重要的初级消费者,其密度对沉水植物及其附着藻类的影响存在争议。本研究设置4种初始螺类密度（0、40、80、240 ind·m^-2）,研究淡水螺类（椭圆萝卜螺Radix swinhoei H.Adams）对刺苦草（Vallisneria spinulosa Yan）及其附着藻类的直接牧食作用和螺类种群的变化。结果显示,在添加螺处理中,刺苦草和人工基质表面附着藻类的生物量显著降低,同时沉水植物的生长显著增加,在较高初始密度螺类处理中刺苦草产生更多的分株。到实验结束时,螺类的死亡率较高,但3个有螺处理间螺类鲜重无显著差异,而高初始密度螺类条件下的最终密度仍较高,同时个体重量（均重）也较小。在中富营养条件下淡水螺类可以直接牧食沉水植物叶片,但对植物生长的抑制作用不显著,有可能是因为沉水植物并不能作为唯一的食物来源维持螺类种群,同时螺类的种群结构受到水体营养水平等因素的制约。     

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.     

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.     

Schistosoma mansoni: influence of Biomphalaria glabrata size on susceptibility to infection and resultant cercarial production

Biomphalaria glabrata snails of the same age, but different sizes, were used to determine size-related susceptibility to Schistosoma mansoni miracidial infection and the influence of snail size on total cercarial production. Snails with shell diameters from less than 5 to greater than 17 mm were individually exposed to one or several miracidia, depending on the experiment. In snails exposed to multiple numbers of miracidia, the percentage of snails which developed patent infections was lower in snails with larger shell sizes. This was also reflected by fewer primary sporocysts per infected snail found in tissues of the larger snails. Upon determining cercarial production in these groups over a 1-month period there were no statistical differences between any groups in the numbers of cercariae produced per snail. However, upon determining the number of successful primary sporocysts found in cohort snails of each size group, cercarial production increased as a function of the number of successful primary sporocysts. This was verified by examining cercarial production in various size snails with known monomiracidial infections. Our data therefore confirm and extend earlier work using snails infected with unknown numbers of miracidia and clearly show that total S. mansoni cercarial development and decreased susceptibility of snails is a direct reflection of snail size and not necessarily age of the snail.