Current velocity and habitat patchiness shape stream herbivore movement

Animal movements are influenced by the structure and arrangement of patches in a landscape. Most movement studies occur in terrestrial landscapes, though aquatic landscapes are equally heterogeneous and feature patches that differ in resistance to animal movements. Furthermore, the variable and highly directional flow of water over streambed landscapes is a unique environmental element, yet its constraint on animal movement is poorly understood. This study examines how habitat availability in a streambed landscape interacts with current velocity to affect movement patterns of two benthic grazers: glossosomatid caddisfly larvae (Agapetus boulderensis) and pulmonate snails (Physa sp.). Using experimental streambed landscapes, we found that Agapetus traveled farther as availability of smooth habitat (composed of low diatom turfs) increased compared to tall, structured filamentous stands, but only did so in slow current velocities. Swifter flows caused restricted movement of Agapetus and more upstream‐oriented paths, but only in smooth landscapes where the potential for flow refugia from filamentous stands was minimal. Similarly, increasing proportions of smooth habitat facilitated greater net displacement of Physa using more upstream‐oriented paths. Higher current velocities caused Physa to move faster, a pattern demonstrated only in smooth landscapes. Our results illustrate a strong interaction between benthic habitat structure and current velocity in shaping patterns of grazer movements in a streambed landscape. Our study also suggests that the flow of water be considered not only a strong environmental gradient in streams, but also an interactive landscape feature that can combine with streambed structure to determine the permeability of patches to the movement of benthic organisms. Landscape ecology has mainly focused on terrestrial environments, and this study offers insight into some of the unique processes that may shape animal movement in aquatic environments.     

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.     

Octopamine boosts snail locomotion: behavioural and cellular analysis

We measured the reduction in locomotion of unrestrained pond snails, Lymnaea stagnalis, subsequent to transdermal application of two selective octopamine antagonists, epinastine and phentolamine. After 3 h in fresh standard snail water following treatment with 4 mM epinastine or 3.5 mM phentolamine, the snails’ speed was reduced to 25 and 56% of the controls (P < 0.001 and P = 0.02, respectively). The snails’ speed decreased as the drug concentration increased. In the isolated CNS, 0.5 mM octopamine increased the firing rate of the pedal A cluster motoneurons, which innervate the cilia of the foot. In normal saline the increase was 26% and in a high magnesium/low calcium saline 22% (P < 0.05 and 0.01, respectively). We conclude that octopamine is likely to modulate snail locomotion, partially through effects on pedal motoneurons.     

Cold tolerance of invasive freshwater snails,Pomacea canaliculata,P. maculata,and their hybrids helps explain their different distributions

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Water velocity preferences of Coho Salmon during the parr-smolt transformation

Juvenile Coho Salmon undergo many physiological changes during their springtime transformation from a freshwater parr to a migratory, seawater-capable smolt. Although field observations indicate smolts moving towards the surface and across the breadth of their streams to either swim or drift downstream with the current, water-velocity preferences of these developing cohos are unknown. Using video analysis of their swimming patterns in a calibrated, laboratory flow table with a velocity gradient, groups of three cohos generally increased their preferred water velocity through the springtime study period, to a late-May peak (daytime data, change-point regression analysis, p < 0.05) and over the entire period (nighttime data, regression analysis, p < 0.05). Moving to swifter currents should facilitate the downstream movements of these young cohos, as they develop through the parr-smolt transformation period. This information should assist managers of regulated watersheds and salmon hatcheries in optimizing juvenile salmon survival (e.g., with timely, late-spring water releases producing 0.1–0.3 m s⁻¹ downstream water velocities).     

Growth and survival of juvenile Pomacea canaliculata (Gastropoda: Ampullariidae) in plain streams associated to different land uses

Pomacea canaliculata is an apple snail that has become an invasive species in several countries. In this research, through two simultaneous experiments, we assessed the effects of different land uses (urban, agriculture, cattle grazing, farms and recreation) on the growth and survival of snails from native populations. The survival was not affected by the water quality of the three streams studied, whereas the growth of snails exposed to the water of the urban stream was impaired. Therefore, our results suggest that the growth rate of P. canaliculata could be useful for the assessment of a moderate urban impact.     

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.     

Coexistence in streams: do source–sink dynamics allow salamanders to persist with fish predators?

Theory suggests that source–sink dynamics can allow coexistence of intraguild predators and prey, but empirical evidence for this coexistence mechanism is limited. We used capture–mark–recapture, genetic methods, and stable isotopes to test whether source–sink dynamics promote coexistence between stream fishes, the intraguild predator, and stream salamanders (Dicamptodon aterrimus), the intraguild prey. Salamander populations from upstream reaches without fish were predicted to maintain or supplement sink populations in downstream reaches with fish. We found instead that downstream reaches with fish were not sinks even though fish consumed salamander larvae—apparent survival, recruitment, and population growth rate did not differ between upstream and downstream reaches. There was also no difference between upstream and downstream reaches in net emigration. We did find that D. aterrimus moved frequently along streams, but believe that this is a response to seasonal habitat changes rather than intraguild predation. Our study provides empirical evidence that local-scale mechanisms are more important than dispersal dynamics to coexistence of streams salamanders and fish. More broadly, it shows the value of empirical data on dispersal and gene flow for distinguishing between local and spatial mechanisms of coexistence.     

Biological control of the exotic invasive snail Pomacea canaliculata with the indigenous medicinal leech Whitmania pigra

The apple snail Pomacea canaliculata has been an invasive species in China for decades and causes enormous losses to agriculture. The predation capability of the leech Whitmania pigra on P. canaliculata was studied for its economic benefit and potential application. In the present study, the leech W. pigra exhibited a strong predatory capacity in controlling P. canaliculata under both laboratory and field conditions, and it showed no bias towards consuming smaller snails during the experiments. More than 80% of the 80 snails (of which half had reached maturity) were preyed upon by 20 developing leeches (2–7?g) in miniature rice fields over a 15-day period, and the number of rice seedlings damaged by P. canaliculata was decreased in the presence of W. pigra. In a separate experiment, 15 developing snails were exposed to caged leeches and conspecific snails for four weeks. The food intake and growth of P. canaliculata were inhibited under the predation risk imposed by W. pigra, but the feeding rate, food conversion efficiency and survival of P. canaliculata were not conspicuously influenced.     

A biomechanical hypothesis explaining upstream movements by the freshwater snail Elimia

1. Many taxa of freshwater invertebrates show active upstream movements, particularly the snails. Hypotheses explaining this behaviour invoke the search for food or space, compensation for drift, avoidance of predation and hydrodynamic effects. The pervasiveness of upstream movements among remote lineages of snails (two subclasses, three orders, 10 families), however, suggests that snails may move upstream for mechanical rather than adaptive reasons.
2. It is proposed that upstream movements by snails are a function of torque on the snail's foot generated by hydrodynamic drag on the shell. When subject to high broadside drag-forces on their shells, snails are able to reduce torque and stabilize orientation only by directing their anterior aspect upstream.
3. Movements of the freshwater pleurocerid snail Elimia were studied by following marked free-ranging individuals in six streams in Alabama, USA (four species, eight populations).
4. Populations showed either no net movement (two streams) or significant upstream movements ranging to a mean of ≈40 m over a 3-month period (four streams). Movement patterns were stream specific rather than species or population specific. Within populations showing significant upstream movements, snails with shell lengths ≤10 mm showed little net movement. Larger snails showed movements from 0 to >200 m upstream.
5. A torque-constrained random walk model was used to perform a post hoc test of the hypothesis that upstream movements were a function of torque on the snail's foot generated by hydrodynamic drag on the shell. The model predicted upstream and size-dependent movement patterns that approximated those observed for snails in the field.     

Diatom communities along stream longitudinal gradients

1. Summer diatom communities on artificial substrates were sampled weekly for a month in three first- to sixth-order tributaries of the Kentucky River to determine how community structure varied with stream size. 2. Diatom cell abundances were generally higher in the headwaters. Species diversity increased in a downstream direction in two of the streams, and in an upstream direction in the third. However, diversity in general seemed more closely related to current regimes than to stream size per se, with highest species diversity at intermediate current velocities. 3. Variation in diatom accumulation rates was greater in downstream communities than in the headwater assemblages of two streams, suggesting that downstream communities may experience greater fluctuations in abundance, at least under low-flow conditions. 4. Patterns of species distributions suggested a relationship between morphological growth forms (guilds) and stream size, as well as the influence of current. Achnanthes spp., Eunotia spp., erect, and stalked taxa were more commonly associated with headwater assemblages. Filamentous and centric diatoms occurred with greater frequency downstream.     

贝类壳-体质量比和静水沉降特性的相关性

软体动物贝壳表型是适应性进化的结果,为探究贝壳形态特性与软体动物随水流扩散功能的相关性,实验选取9种具代表性的腹足纲和双壳纲贝类作为研究对象,研究了其成体的壳-体质量比与静水沉降特性及二者的相关性。结果表明：体质量的变异系数（C.V.=1.11）大于壳长（C.V.=0.67）、壳宽（C.V.=0.54）的变异系数;壳-体质量比具较强的种属特异性,种内变异系数小（C.V.=0.20）,而种间差异极显著（P < 0.01）。受试贝类的平均壳-体质量比为0.32±0.13,陆生贝类的壳-体质量比显著小于水生贝类（P < 0.01）,壳-体质量比最大的河蚬（0.54±0.06）是最小的非洲大蜗牛（0.17±0.04）的3倍,福寿螺的壳-体质量比为0.50±0.06。双壳类和螺类的沉降行为差异明显,只有中华园田螺、铜锈环棱螺和河蚬不能在静水中漂浮;沉降速度最大的河蚬（（24.99±4.22）cm/s）是最小的椭圆萝卜螺（（4.13±0.96）cm/s）的6倍,入侵种非洲大蜗牛（（18.30±3.64）cm/s）和福寿螺（（21.77±5.23）cm/s）与土著种铜锈环棱螺（（19.48±3.14）cm/s）和中华园田螺（（21.44±3.92）cm/s）在沉降速度上无显著差异（P < 0.01）。成体贝类的壳-体质量比与沉降速度（R²=0.28）、沉降加速度（R²=0.39）之间存在一定的相关性。在此基础上,贝类沉降特性随生活史变化的研究将进一步揭示贝类进化与种群扩散中的生态水力学作用。     

Secondary production and diet of an invasive snail in freshwater wetlands: implications for resource utilization and competition

Invasive species can monopolize resources and thus dominate ecosystem production. In this study we estimated secondary production and diet of four populations of Pomacea canaliculata, a freshwater invasive snail, in wetlands (abandoned paddy, oxbow pond, drainage channel, and river meander) in monsoonal Hong Kong (lat. 22°N). Apple snail secondary production (ash-free dry mass [AFDM]) ranged from 165.9 to 233.3 g m⁻² year⁻¹, and varied between seasons. Production was lower during the cool dry northeast monsoon, when water temperatures might have limited growth, but fast growth and recruitment of multiple cohorts were possible throughout much (7–10 months) of the year and especially during the warm, wet southwest monsoon. The diet, as revealed by stomach-content analysis, consisted mainly of detritus and macrophytes, and was broadly consistent among habitats despite considerable variation in the composition and cover of aquatic plants. Apple snail annual production was >10 times greater than production estimates for other benthic macroinvertebrates in Hong Kong (range 0.004–15 g AFDM m⁻² year⁻¹, n = 29). Furthermore, annual production estimates for three apple snail populations (i.e. >230 g AFDM m⁻² year⁻¹) were greater than published estimates for any other freshwater snails (range 0.002–194 g AFDM m⁻² year⁻¹, n = 33), regardless of climatic regime or habitat type. High production by P. canaliculata in Hong Kong was attributable to the topical climate (annual mean ~24°C), permitting rapid growth and repeated reproduction, together with dietary flexibility including an ability to consume a range of macrophytes. If invasive P. canaliculata can monopolize food resources, its high productivity indicates potential for competition with other macroinvertebrate primary consumers. Manipulative experiments will be needed to quantify these impacts on biodiversity and ecosystem function in wetlands, combined with management strategies to prevent further range extension by P. canaliculata.     

Field observations of the alarm response to crushed conspecifics in the freshwater snail Pomacea canaliculata: effects of habitat,vegetation, and body size

The freshwater snail Pomacea canaliculata shows alarm responses to chemical cues released from injured conspecifics, but its behavioural responses in the field are unknown. We investigated effects of habitat (canals or paddy fields), vegetation, and body size on alarm responses in the field. Snails responded to crushed conspecifics within 4 min by burying themselves, but the proportions of self-buried snails were generally lower (0–28% depending on experiments) than those reported in the laboratory. Snails not only showed the alarm response, but also frequently fed on crushed conspecifics. There were no influences of habitat or body size on the proportions of individuals showing the alarm response. Nevertheless, in paddy fields with high-density vegetation a higher proportion of snails showed the alarm response than in paddy fields with low-density vegetation.     

Changes in chemical components in the freshwater apple snail, Pomacea canaliculata (Gastropoda: Ampullariidae), in relation to the development of its cold hardiness

The apple snail, Pomacea canaliculata, is an invasive freshwater snail. It increases its cold hardiness before winter. However, the physiological mechanism of cold hardiness in molluscs is poorly understood, especially in freshwater molluscs. In this study, we examined the changes in low molecular weight compounds, glycogen and lipids, in the body of P. canaliculata in association with the development of cold hardiness. When snails without cold hardiness were experimentally cold-acclimated, the amount of glycerol, glutamine, and carnosine increased, while glycogen and phenylalanine decreased. Overwintering cold-tolerant snails collected from a drained paddy field in November also showed increased glycerol in their bodies with decreasing glycogen concentration, compared to summer snails collected from a submerged field. Water content also decreased during the cold acclimation, although the water loss was minimal. These results indicate that the freshwater snail, P. canaliculata enhances cold hardiness by accumulation of some kinds of low molecular weight compounds in its body as some insects do. However, the actual function of each low molecular compound is still unknown.     

Velocity and turbulence distribution around lotic macrophytes

Flow velocity and turbulence patterns were measured in and around a common lotic macrophyte, Ranunculus penicillatus subspecies pseudofluitans (stream water-crowfoot), using a two-dimensional electromagnetic current meter (EMCM). Due to the high shooting density of this species, there was a sharp velocity gradient at the plant boundary, with velocities dropping to a constantly low value after no more than 5 cm into the plant, thus forcing most of the flow over and around the macrophyte. There was a dead-water zone immediately downstream of the plant, beyond which the current moved in from the sides to allow flow under the trailing shoots. High turbulence intensities were recorded for both downstream and cross-stream velocity components at the lateral margins and downstream of the plant. Meanwhile, pulses of water upstream of the plant produced turbulence in the downstream component, but not in the cross-stream component.     

Advertising acceptability: is mollusk olfaction important in seedling selection?

Although seedling herbivory is an important selective filter in many plant communities, how and why seedlings are selected is poorly understood. Here, we examined the putative role of herbivore olfaction in dictating seedling selection. Using a Y-tube olfactometer we compared snail (Helix aspersa) preference for pellets derived from 14-day-old macerated seedlings of nine European grassland (‘Test’) species against standard (‘Control’) pellets derived from lettuce. Snail movement towards ‘Test’ pellets was strongly correlated with seedling acceptability (Pearson’s r ² = 0.86, P > 0.01) and where snails exhibited a positive choice for the ‘Test’ species, the choice was made more quickly for highly acceptable species (r ² = 0.86, P > 0.01). In elucidating a link between seedling acceptability and olfactory response to macerated seedlings, our study suggests that even from an early ontogenetic stage plant selection by snails may be governed by olfactory cues. This finding highlights the need for research on the role of plant volatiles in plant–herbivore interactions to consider more fully interactions operating at the seedling stage.     

A preliminary investigation into the use of the invasive golden apple snail,Pomacea canaliculata (Lamarck, 1822), as a water purifier and food source in the breeding ponds of the oriental weatherloach Misgurnus anguillicaudatus (Cantor, 1842)

This study investigated the feasibility of using an invasive snail, Pomacea canaliculata, as a food source and water purifier for the commercial breeding of the loach Misgurnus anguillicaudatus. The predatory potential of M. anguillicaudatus (3.5–5.5?g) against hatchling snails was evaluated in aquaria and simulated paddy fields. Some hatchling snails left the water to avoid being preyed upon by the loaches, and approximately 10 hatchlings died per day in the presence of five loaches in aquaria, whereas a weaker snail control effect was observed in the simulated paddy fields. The growth of rice seedlings (Oryza sativa) was not reduced by the presence of hatchling snails alone, but the shoot biomass of seedlings coexisting with snails was promoted after introducing the loaches. Additionally, the presence of P. canaliculata adults improved the aquatic environment in the short term for loach breeding by decreasing the turbidity of the water. Importantly, M. anguillicaudatus (12–18?g) mortality decreased and its weight increased in the presence of adult snails.     

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     

Dispersal and upstream migration of an amphidromous neritid snail: implications for restoring migratory pathways in tropical streams

1. The amphidromous life cycle of several species of neritid snails, shrimp and gobies throughout the tropics includes juveniles that migrate from the ocean to breed in fresh water. In many Hawaiian streams, the decline of Neritina granosa, an endemic gastropod, has been associated with habitat degradation and water withdrawal, which are common factors affecting tropical rivers around the world. 2. We investigated the effects of water withdrawal and density on dispersal and upstream migration of N. granosa using three experimental treatments: (i) reduced flow (RF) owing to a stream diversion, (ii) natural flow (NF) and (iii) natural flow with artificially increased snail density. For each treatment, snails were differentially tagged and released in a stream without a natural, extant population of N. granosa. 3. Capture rates ranged from 17 to 65% over a 63‐day period following release. Captures on 2–6 days after release measured initial dispersal and migration, whereas longer‐term migration rates were calculated from snails captured 16–63 days after release. Snails under NF displayed positive rheotactic behaviour, with only 3–12% demonstrating initial downstream movement. Under RF, 22–77% of snails moved downstream or showed no bias either way. 4. Initial mean upstream migration rate (UMR) was 0.25, 0.66 and 1.16 m day^?1 under RF, NF and natural flow with increased snail density, respectively. Longer‐term migration rates did not differ significantly between treatments, and the overall mean was 0.62 m day^?1. 5. Principal component analysis and generalised linear models were used to identify habitat characteristics important to UMR, with habitat and reach‐scale hydraulics as the most important factors. The relationship between discharge and UMR suggested it would take 11–35 years for snails to migrate past the most upstream water diversion. However, rates from published studies of neritid snail species migrating en masse or in long lines under natural situations suggested that N. granosa could migrate above stream diversions within 72 days–2.5 years (when in an aggregation) and 29 days–1.1 years (when following in long lines of individuals). 6. An understanding of upstream neritid snail migration can be used for the management and conservation of this and other migratory species in tropical streams.