A Population Growth Trend Analysis for Neotricula aperta,the Snail Intermediate Host of Schistosoma mekongi,after Construction of the Pak-Mun Dam

Background

The Pak-Mun dam is a controversial hydro-power project on the Mun River in Northeast Thailand. The dam is sited in a habitat of the freshwater snail Neotricula aperta, which is the intermediate host for the parasitic blood-fluke Schistosoma mekongi causing Mekong schistosomiasis in humans in Cambodia and Laos. Few data are available which can be used to assess the effects of water resource development on N. aperta. The aim of this study was to obtain data and to analyze the possible impact of the dam on N. aperta population growth.

Methodology/Principal Findings

Estimated population densities were recorded for an N. aperta population in the Mun River 27 km upstream of Pak-Mun, from 1990 to 2011. The Pak-Mul dam began to operate in 1994. Population growth was modeled using a linear mixed model expression of a modified Gompertz stochastic state-space exponential growth model. The N. aperta population was found to be quite stable, with the estimated growth parameter not significantly different from zero. Nevertheless, some marked changes in snail population density were observed which were coincident with changes in dam operation policy.

Conclusions/Significance

The study found that there has been no marked increase in N. aperta population growth following operation of the Pak-Mun dam. The analysis did indicate a large and statistically significant increase in population density immediately after the dam came into operation; however, this increase was not persistent. The study has provided the first vital baseline data on N. aperta population behavior near to the Pak-Mun dam and suggests that the operation policy of the dam may have an impact on snail population density. Nevertheless, additional studies are required for other N. aperta populations in the Mun River and for an extended time series, to confirm or refine the findings of this work.     

UPTAKE OF ACETATE BY NEOTRICULA APERTA (GASTROPODA: POMATIOPSIDAE), THE SNAIL HOST OF SCHISTOSOMA MEKONGI IN THE LOWER MEKONG BASIN

All three races of Neotricula aperta, an epilithic, schistosometransmitting, snail of the Mekong and Mul rivers of NortheastThailand and southern Laos, were found to take up acetate froma dilute solution. After 48 h incubation the mean specific netuptake rates (µmol^–1 g^–1 h^–1), from750 µM acetate, were: 1.86, -race; 1.39, ß-raceand 3.25, y-race. Over 48 h the snails were able to achievereductions in the ambient acetate concentration of up to 60%.Incubations under bacteriostatis suggested that bacteria arenot involved in the uptake of acetate by N. aperta. The uptakecharacteristics conform to the Michaelis-Menten model. The transportconstants, J_max (µmol^–1 g^–1 h^–1) andK_t (µM) were 1.10 and 102 respectively (-race). Racialdifferences in uptake characteristics are discussed in relationto micro-habitat differences. HPLC indicated concentrations of acetate in y-N. aperta microhabitatsof around 325 µM. This suggests a pool size sufficientto satisfy only 6% of the snail's basal metabolic rate (BMR).Levels within the epilithic aufwuchs, however, are probablyhigh enough to provide for more than 50% of the BMR. The possible role of acetate in the energy metabolism of N.aperta is discussed. Short-chain carboxylic acids (such as acetate),arising from the decomposition of the aufwuchs, could representsources of fermentable organics that may be taken up by N. apertasnails and used to supplement their nutrition during times offood shortage. However, further investigations involving ¹⁴C-labellingtechniques are required. The findings of this investigationhave implications for the chemical ecology and life-cycle ofN. aperta. (Received 16 June 1994; accepted 28 July 1994)     

POPULATION DENSITY EFFECTS ON GROWTH IN CULTURE OF THE EDIBLE SNAIL HELIX ASPERSA VAR. MAXIMA

The effects of population density on the growth of H. aspersaMÜller var. maxima under controlled environmental conditionswere examined. Inhibitory effects on snail growth and maturityresulting from increased population density, between 100–800snails m^–2 of floor area, were observed for a range ofcontainer cleaning frequencies. At all population densities,enhanced snail growth was observed when the frequency of containercleaning was increased to a two-day interval. No significantdifferences were recorded, following 19 weeks growth, betweenfinal mean weights of snails from containers cleaned less frequently.The lowest snail mortality was consistently recorded at thelowest population density in the most frequently cleaned containers.At all snail population densities three phases of growth wereobserved: (a) lag (0–5 weeks), (b) rapid (6–15 weeks)and (c) stable (16 weeks and over). During the first three weeksof growth, high population density had a positive effect ondiet consumption, food conversion efficiency and snail growth.Adverse population density effects increased progressively duringphase (b), typically following 9 weeks growth. Juvenile snailstransferred from high to low population densities during phase(b) continued to exhibit slower growth rates associated withhigh population densities. Food conversion efficiency of snailsin all treatments decreased throughout the experimental periodbut with no overall effect of container cleaning frequency apparent.Inherent growth variability of sibling snails was unaffectedby population density or container cleaning frequency. The importanceof the results for intensive snail culture is discussed. (Received 23 June 1994; accepted 1 December 1994)     

BIOCHEMICAL COMPOSITION OF HELIX SNAILS: INFLUENCE OF GENETIC AND PHYSIOLOGICAL FACTORS

This paper describes the biochemical composition of differentspecies (Helix lucorum, Helix pomatia) and sub-species of snails(Helix aspersa aspersa, Helix aspersa maxima) reared in thesame conditions with a feed (‘Helixal’) speciallydesigned for edible snails. In addition, the composition ofwild H. pomatia and H. lucorum is presented to allow comparisonbetween snails of different origins. Analyses determined thepercentages of proteins, lipids and minerals. They reveal bothsimilarities and differences in composition according to thespecies and the part analysed (whole body, pedal mass, and visceralmass). H. pomatia contains the highest percentage of mineralmatter and the lowest percentage of lipids. Surprisingly, proteincontents are slightly different between artificially rearedH. aspersa maxima of 3 months old and wild H. pomatia. The resultsmake it possible to evaluate nutritional quality of snails withthe composition of the body of four edible snail species. (Received 16 September 1996; accepted 24 May 1997)     

TAXONOMIC REVIEW OF THE TREE SNAIL AMPHIDROMUS ALBERS, 1850 (PULMONATA: CAMAENIDAE) IN THAILAND AND ADJACENT AREAS: SUBGENUS AMPHIDROMUS

Extensive series of Amphidromus Albers, 1850 were collectedfrom low-altitude areas (30–500 m above sea level)in Thailand, Laos, Malaysia, Singapore and Indonesia between1993 and 2004. Data on shell colour, genital and spermatophoremorphology, and radular structure are presented. Five speciesof Amphidromus (Amphidromus) are reviewed, of which four occurin Thailand. Intraspecific variation between populations ofAmphidromus (Amphidromus) atricallosus (Gould, 1843) is considered;three subspecies are accepted and a new subspecies described.Subspecies of A. (A.) inversus (Müller, 1774) and A. (A.)schomburgki (Pfeiffer, 1861) are reconsidered, and a new subspeciesof the latter is described. A dichotomous key to the speciesand subspecies of Amphidromus (Amphidromus) in Thailand is provided. (Received 7 May 2004; accepted 31 March 2005)     

THE RELATIONSHIP BETWEEN ENVIRONMENTAL VARIABLES AND THE DENSITY OF THE MUD SNAIL HYDROBIA TOTTENI IN A NOVA SCOTIA SALT MARSH

The relationship between environmental variables and the densityof Hydrobia totteni was examined in a restricted area of LawrencetownMarsh, Nova Scotia. Densities averaged 50/m² in Spartina alterniflora,9800/m² in Zostera marina, and 19,000/m² in sand. Of 8 environmentalfactors considered, depth and sediment grain size were the mostimportant in explaining variations in snail density. Since thesefactors together explained only 42.0% of the total variationno single physical factor controls the density of Hydrobia ina restricted area, though salinity does control densities overa wide area. Salinity and temperature tolerances and the abilityof the snails to select appropriate sediment grain sizes weredetermined experimentally and related to the distribution ofthe snails in the marsh. ^*Present address: Western Australian Museum, Francis Street,Perth, Western Australia, 6000 (Received 20 June 1976;     

FIELD OBSERVATIONS ON FEEDING OF THE LAND SNAIL HELIX ASPERSA MULLER

Feeding of the land snail Helix aspersa (Müller) was observedat monthly intervals. Three natural populations in Galicia (NW-Spain)were studied. At two sites only a few plants constituted thebulk of the diet and in spring the snails' diet had the highestdiversity (H'). In the third population feeding and distributionof Helix aspersa (Müller) were observed in a small plotwith permanent patches of Urtica dioica. Nearly one half offeeding snails fed upon Urtica dioica. Most of the other observations wereon Mentha suaveolens, Ranunculus repens and Gramineae. The diversity ofthe snails' diet showed seasonal variation with the maximumin the autumn months. Comparison between the availability ofthe different plant species and their contribution to the snails'diet showed that the snails did not eat at random; Urtica dioicawas eaten much more than expected from its occurrence and grasseswere strongly under-represented in the snails' diet. Temporalchanges of availability were significantly correlated with the amountseaten in the case of Urtica, but not for the other food plants.The distribution of the snails in the plot was significantlycorrelated with that of Urtica. Chemical analyses of the foodplants revealed Urtica as the species with the higher protein,ash and calcium contents. The strong preference of Helix aspersafor Urtica dioica could be explained by the value of Urticaas food or by its suitability as habitat for the snails. The largestproportions of green material in the snails' diet occurred inthe spring and juveniles ate more green material than adultsin the three populations. (Received 16 March 1998; accepted 30 November 1998)     

HETEROGENIC COPULATORY BEHAVIOUR PRODUCES NON-RANDOM MATING IN LABORATORY TRIALS IN THE LAND SNAIL HELIX ASPERSA MULLER

This study reports the results of mate-choice experiments betweenresident and non-resident individuals of the simultaneous hermaphonteland snail Helix aspersa. Snails from different sites differedin their mating proclivity, which resulted in non-random matingin laboratory mate-choice tests. Those snails with the highermating propensity therefore tended to mate with their own typeand tended to be chosen as partner because they were also activelyseeking mates. Keeping snails under crowded conditions did noteffect a lasting influence on mating behaviour in laboratorytests, despite the decline in copulatory behaviour under crowdedfield conditions. (Received 7 November 1994; accepted 31 August 1995)     

INVESTIGATIONS OF THE ROLE OF MUCUS AND FAECES IN INTERSPECIFIC INTERACTIONS OF TWO LAND SNAILS

A previous report (Smallridge & Kirby, 1988) showed interspecificcompetition between the two land snails Theba pisana and Cernuellavirgata and suggested the mechanism of competition may be thatheterospecific mucus and excrement inhibits activity. We describea series of experiments in which we could find no consistentevidence for any short-term inhibition of activity in the presenceof heterospecific mucus. Furthermore snails did not consistentlyavoid mucus-treated surfaces or mucus-coated food. Cernuellastopped at and fed from snail faeces (both conspecific and heterospecific)more than did Theba. This suggests an alternate mechanism forinterspecific competition. (Received 9 April 1992; accepted 9 June 1992)     

FOOD PREFERENCES AND DIETARY OVERLAP BY TERRESTRIAL SNAILS IN LOGOS AREA (EDESSA, MACEDONIA, NORTHERN GREECE)

Among the five coexisting species of snail in Edessa (N. Greece),Bradybaena fruticum and Helix lucorum dominated in density andbiomass in comparison with Xeropicta arenosa, Monacha cartusianaand Cepaea vindobonensis. Resting places of each species inrelation to plant cover and diet preferences are examined seasonallyand throughout a year. Discriminant analysis for the annualdata showed that 83% of the snail group species could be differentiatedby the plants on which they were attached. Each snail specieswas usually found sitting on one of the most abundant plants.Cepaea preferred to sit on and eat senescent material even inspring time and Bradybaena green material. In spring, the snailsdid not consume plant material in proportion to the occurrenceof those plants but they usually did so in summer and in autumn.Hedera helix was not consumed even when it was abundant. Inspring and autumn the snails had consumed plants to which theywere found attached, but in summer they did not. (Received 9 August 1993; accepted 14 February 1994)     

DISTRIBUTION AND PRODUCTION OF THREE HYDROBIA SPECIES (GASTROPODA: HYDROBIIDAE) IN A SHALLOW COASTAL LAGOON IN THE BAY OF CADIZ, SPAIN

The abundance, life span, growth and production of the mud snailsHydrobia minoricensis, H. ulvae and H. ventrosa in a semi-naturallagoon system were studied by taking monthly samples at threesites during 1991 and 1992. The most abundant species, H. minoricensisoccurred at mean densities of 12834 to 26264 snails m^–2(10.7 to 25.8g dry weigh m^–2), depending on the site.The least abundant species, H. ulvae, occurred at mean densitiesof 185 to 353 snails m^–2 (3.2 to 2.2g dry weight m^–2).The numerical abundance and biomass of the three Hydrobia specieswere positively related to the biomass of benthic macroalgae(P<0.01). Although H. ulvae egg capsules were recorded throughoutthe year, newly hatched snailsof this species were not observed,in contrast to the other two species. The early spring and summercohorts of H. minoricensis and H. ventrosa seemed to be themost numerous. The average life spans of these two species wereestimated to be about 18 and 13 months respectively. Annualproduction estimates for the whole lagoon system were 29.0 (6.3),5.5 (0.8) and 5.2 (1.0)g dry weight (ash-free dry weight) m^–2yr^–1 for H. minoricensis, H. ulvae and H. ventrosa respectively.The annual P/B ratio was about 2 for H. minoricensis and H.ventrosa. (Received 5 July 1994; accepted 5 October 1994)     

WATER EXCHANGE, TEMPERATURE TOLERANCE, OXYGEN CONSUMPTION AND ACTIVITY OF THE NAMIB DESERT SNAIL, TRIGONEPHRUS SP.

Water exchange, temperature tolerance and oxygen consumptionof the snail, Trigonephrus sp., from the southern Namib desertof Namibia were examined and related to activity. At 25°Cand 15% R.H. mean water loss and food and water uptake were5.95 mg. day^–1 and 630 mg.day^–1, respectively. Bodytemperature tracked sand temperature. Snails tolerated sandtemperatures as high as 45°C. Mean ± S.D. oxygenconsumption rates were 32.0 ± 2.94 µlO₂.g totalbody mass^–1.h^–1 at 15°C, when the snails wereactive, and 11.27 µlO₂.g total body mass^–1.h^–1at 25°C, when the snails were inactive. These values are2-6 times lower than those recorded for the similarly sizedmesic snail, Helix aspersa. Activity experiments indicated thatlow ambient temperatures and high humidities were favoured bythe snails. This, together with the burying behaviour of thesesnails during high temperatures, suggests that they limit stressby restricting activity to physiologically-favourable periods,even though more-extreme conditions may be tolerated. (Received 7 June 1990; accepted 20 November 1990)     

CONTRIBUTION OF GENETIC AND ENVIRONMENTAL FACTORS TO SHELL SHAPE VARIATION IN THE LOTIC SNAIL SEMISULCOSPIRA REINIANA (PROSOBRANCHIA: PLEUROCERIDAE)

The genetic and environmental factors affecting shell shapein the freshwater snail Semisulcospira reiniana at Takahashiin Kyoto, central Japan, were studied by means of a rearingexperiment and field observations. Shell shape was characterizedby three parameters; W (whorl expansion rate), T (whorl translationrate), and S (roundness of generating curve). Estimated heritabilitieswere low in all three parameters and the largest component ofthe great shell variation in the Takahashi population was environmentalvariance, suggesting that the main source of shell variationwas phenotypic modulation, in response to the external environment.In the field, snails that were active in fast currents had largerW and smaller T, that is, a larger body whorl and a lower spire,than snails in slow currents. Substratum conditions relatedto T in resting periods although its cause was unclear. Mechanismsare suggested which in the absence of selection of genetic variationcould cause and maintain shell variation in S. reiniana in differentmicrohabitats. (Received 8 March 1996; accepted 3 November 1997)     

EFFECTS OF HEAVY METALS ON SURVIVAL AND GROWTH OF BIOMPHALARIA GLABRATA SAY (GASTROPODA: PULMONATA) AND INTERACTION WITH SCHISTOSOME INFECTION

The effects of dissolved Pb, Cd and Hg on growth and survivalof adult Biomphalaria glabrata Say. uninfected and infectedwith Schistosoma mansoni Sambon, were examined. Pb at concentrationsfrom 0.25–100 µM, Cd from 0.075–0.25 µMand Hg from 0.25–1 µM significantly reduced growthand survival of normal snails. With each metal the effects increasedwith increasing concentration. The LC 25 at 2 wks exposure was82 ± 19, 0.22 ± 0.04 and 0.94 ± 0.13 µM(x ± S.E.) for Pb, Cd and Hg, respectively. Snails exposedto heavy metals continued to be reproductively active, but theegg production was highly variable and no significant effectof heavy metal exposure was demonstrated. Infection also decreasedsurvival and growth of snail hosts and infected individualsexposed to heavy metals displayed the greatest mortality. TheLC 25 for infected snails at 2 wks exposure was 8 ± 3,0.9 ± 0.02 and 0.29 ± 0.08 for Pb, Cd and Hg respectively.A significant interaction between heavy metal exposure and infectionwas apparent. Infected snails were not reproductively active.Cercarial shedding by infected snails was significantly reducedin the presence of heavy metals and by 6 wks shedding had ceasedat the highest metal concentrations. (Received 20 May 1996; accepted 31 July 1996)     

Aaptosyax grypus,a new genus and species of large piscivorous cyprinids from the middle Mekong River

A new genus and species of cyprinid fishes,Aaptosyax grypus, is described from the middle Mekong River of Laos and Thailand, This large and rarely seen species reaches a full meter in length and lives in the main stem of the river. The species is a piscivore and the southernmost representative of the aspiin lineage. Among the aspiins it is unique in having a well-developed adipose eyelid and large symphyseal knob on the mandible which fits into a deep notch in the upper jaw.     

ASPECTS OF LIFE CYCLE, POPULATION DYNAMICS, GROWTH AND SECONDARY PRODUCTION OF THE PULMONATE SNAIL CEPAEA VINDOBONENSIS (FERUSSAC, 1821) IN NORTHERN GREECE

The life cycle, population dynamics, growth and secondary productionof the land snail C. vindobonensis were studied in northernGreece. Demographic analysis of the populations of C. vindobonensisrevealed that a) three cohorts were present in the field throughoutthe year, b) the reproductive period started in late April-Mayand the newly hatched snails appeared in the beginning of June,and c) increased growth rates were observed during spring andearly summer, but also during autumn for the newly hatched snails. According to von Bertalanffy's method C. vindobonensis needs7 years to attain its maximum size measured in the field. Mortalityrate is very high during the first year of life, while lifeexpectancy is higher during the second year of life and decreasesafterwards. Net reproductive rate (R_o) was equal to 3.1 andthe finite capacity for increase (antilog_er_c) was equal to 1. Estimated annual secondary production with Hynes' frequencymethod revealed a mean standing crop (B) of 0.99 g/m²/year anda production (P) of 1.3 ± 0.11 g/m²/year. Annual turnoverratio (P/B) was equal to 1.31. (Received 7 April 1997; accepted 2 October 1997)     

COPULATORY ACTIVITY AND SPERM PRODUCTION IN BULINUS (PHYSOPSIS) GLOBOSUS (GASTROPODA: PLANORBIDAE)

Young Bulinus (Physopsis) globosus performed male copulatoryactivity and cross-fertilized other snails before their femalereproductive tracts were mature. The two most immature snailsshowed preputial eversion when secretion was present only inthe muciparous gland and at the carrefour region of the oviduct.Sixteen snails showed preputial eversion and four snails cross-fertilizedother snails when their oothecal glands and/or major portionof the oviducts contained either no secretion or only scantyamounts. When paired with a partner snail for 12 or 20 consecutive days,adult snails copulated as males on approximately 60% of thedays paired and up to 8 consecutive days. Virgin snails raisedin isolation copulated as male at the same rate as non-virgin,community-raised snails. Ability to copulate as male was notdependent upon previous experience as male or female. Aftera single copulation as male after 7 days isolation, the hermaphroditicducts of maleacting snails contained 87 000 sperm. Sperm productionoccurred at approximately 50 000 sperm.d-¹, until at 10 dayspost-copulation, snails contained 639 000 sperm. (Received 25 May 1982;     

LIFE HISTORY AND POPULATION DYNAMICS OF COCHLICELLA ACUTA (MuLLER) (GASTROPODA: HELICIDAE) IN A PASTURECEREAL ROTATION

The conical snail Cochlicella acuta was sampled over a 3-yearperiod in a pasture-cereal rotation and a nearby roadside wastelandat Hardwicke Bay, South Australia. The life cycle of C. acuta was primarily biennial in the agriculturalfields, with offspring being produced in large numbers in thepasture phase but not the cereal phase of the rotation. Thebreeding season lasted from autumn to spring. Snails were mostabundant in spring and summer, especially near the edges offields. During summer, snails aggregated on robust weeds suchas Reseda lutea. Many snails were killed by burning pasturesin autumn, prior to sowing crops. Snails were more abundantand smaller in size in the roadside wasteland than in the agriculturalfields. Strategies for the control of C. acuta in pasture-cereal rotationsare discussed. (Received 9 April 1990; accepted 10 August 1990)     

OBSERVATIONS ON THE ECOLOGY OF BULINUS (PHYSOPSIS) GLOBOSUS (MORELET), B. (BULINUS) FORSKALII (EHRENBERG) AND LYMNAEA NATALENSIS (KRAUSS) IN RICE SWAMPS IN SIERRA LEONE.

Changes in population of Bulinus globosus, B. forskalii andLymnaea natalensis were studied in four different locationsnamed Sites A, B, C. and D in Bo town Sierra Leone. The snailpopulations and oxygen concentrations of water increased inthe rainy season. The biological oxygen demand did not havesignificant correlation with changes in snail populations inSites B and D. B. globosus was absent from Site A while B. forskaliiwas the predominant snail in Site C. Site D with the highestmean monthly conductivity of 176 µmhos contained 54% whileSite A with a similar reading of 90µmhos contained 1%of all snails found. (Received 22 May 1980;     

COPULATION AS FEMALES AND USE OF ALLOSPERM IN THE FRESHWATER SNAIL GENUS BULINUS (GASTROPODA:PLANORBIDAE)

Each of 8 snails in 2 groups of Bulinus (Physopsis) globosus,1 group raised in isolation and 1 group raised in community,were paired for 14 consecutive days with a male-acting partnersnail. In each group, the experimental snails, which were notallowed to act as males, were able to copulate as females onapproximately 94% of the days paired. Two copulations as female,with the same male partner, occurred on 50% of the days thatthe snails were paired, in the 2 groups combined. Non-receptivefemale behaviour by the experimental snails occurred frequently,and copulation was prevented by such behaviour during 6 pairings,3 in each group. Young B. (P.) africanus first copulated as females when theywere 31–33 days old. The accessory sex glands of the femalereproductive tracts of these young female-acting snails containedmoderate to large amounts of secretion. B. (P.) africanus, which were raised in pairs, laidcross-fertilizedeggs in isolation for an average of 76 days, and 1559 eggs/snailwere deposited before cross-fertilization ceased. Cross-fertilizedeggs were produced for as long as 120 days. After 1 copulation as female, virgin B. (P.) africanus laidcross-fertilized eggs for an average of 78 days and deposited3654 eggs/snail before crossfertilization ceased. Cross-fertilizedeggs were produced for as long as 113 days. After 2 copulationsas female, 1 copulation on each of 2 consecutive days, virginB. (P.) africanus laid cross-fertilized eggs for an averageof 102 days and produced 4397 eggs/snail before cross-fertilizationceased. Cross-fertilized eggs were produced for as long as 123days. Snails which were homozygous for an allele governing mantlepigment pattern were raised with a partner which was homozygousfor a different pigment pattern. Young produced in a 4-day periodafter the snails were isolated were 100% heterozygous. The snailswere then rearranged into pairs with a partner of the same genotypefor 4 days, during which time 26% of the young produced werehomozygous. The snails were again isolated for 4 days, and 49%of the young produced during this 4-day period were homozygous.The results of this experiment strongly suggest that multipleoutcrossing occurred. In B. (P.) africanus, stored allosperm were used to fertilizeeggs after 1, 4 and 7 weeks of starvation; after 1 and 4 weeksof 15°C low temperature and 4 weeks of 15°C + 4 weeksof 10°C low temperature; and after 1 and 4 weeks of desiccation.After 8 weeks of desiccation, 2 of 3 surviving snails reproducedby self-fertilization and 1 snail did not reproduce. Too fewsnails survived 8 weeks of desiccation for a conclusion to bereached on the ability of allosperm to survive. (Received 1 June 1984;