Mating behavior,egg laying,and embryonic development in the South American apple snail Asolene pulchella (Ampullariidae,Caenogastropoda)

Asolene pulchella is a dioecious freshwater snail from the La Plata basin, belonging to the Ampullariidae family. Our aim was to study the reproductive biology, including mating behavior, egg laying, and embryonic development under laboratory conditions. Copulations occurred underwater and lasted 2.66?h on average. The males produce fusiform paraspermatic cells (with seven to nine flagella) in addition to euspermatic cells with a corkscrew head, straight middle piece and long flagellum. Females stored sperm for up to 169?days. The egg masses (containing 98.1 eggs on average) are deposited underwater and are composed of many gelatinous packs of 1–4 eggs; the jelly matrix presented numerous calcite microcrystals. The spheroidal egg capsules measured 2.25?mm and contained a yellowish perivitellus of rubbery consistence. The embryonic development extends for 10.8?days on average. Hatchlings measured 1.4?mm and remained on the egg mass, feeding on the jelly. They resemble miniature adults but only began to feed on senescent plant material in their third week of age and begin aerial lung respiration after 8–10?weeks. The details of the structure of the egg masses and development constitute a rich source of traits for comparative and taxonomic studies in the Ampullariidae.     

Inheritance of longitudinal shell bands in the snails <Emphasis Type="Italic">Littorina obtusata</Emphasis> and <Emphasis Type="Italic">L. saxatilis</Emphasis> (Gastropoda,Prosobranchia)

The hypothesis of a monogenic inheritance of dark longitudinal bands on the shell in the gastropods Littorina obtusata and L. saxatilis was checked. One gene having two alleles proved to be responsible for the shell banding pattern in both of the species. The presence of bands was a dominant character in either case.     

Inheritance of longitudinal white shell bands in the snail <Emphasis Type="Italic">Littorina obtusata</Emphasis> (Gastropoda,Prosobranchia)

The hypothesis on monogenic inheritance of white longitudinal shell bands in the snail Littorina obtusata was tested. The data showed that a single gene with two alleles was responsible for the presence of the bands on the snail shell. The presence of bands was the dominant character.     

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     

Species identification from archived snail shells via genetic analysis: a method for DNA extraction from empty shells

During the summer of 2009, apple snails (Mollusca: Ampullariidae) of the genus Pomacea were found to have established a self-sustaining population for the first time in Europe (Ebro Delta, Catalonia, Spain). This group is known for its genetic diversity and at least two distinct species are represented in a clade typified by Pomacea canaliculata. Various citations refer to egg size and color, and clutch size, in addition to shell morphology, for distinction between species. However, due to overlapping characters and phenotypic plasticity, genetic analyses are becoming more widely used methods for this group. Herein, we present a method for DNA extraction to identify species with sufficient precision from empty shells recovered from the field. This technique may be of use more widely for analyses of shells archived in museum collections for phylogenetic and taxonomic studies.     

Preliminary observation on the role of conditioning and mechanical Interference with egg masses and Juveniles in the competitive relationships Between Helisoma Durvi (Wetherby) and the Intermediate host of Schistosoma Mansoni Sambon: Biomphalaria Camerunensis (Boettger).

Biological control of the intermediate hosts of schistosomiasis by means of the introduction oa a competitor snail Helisoma duryi has been suggested. The effect of conditioning by Biomphalaria camerunensis and H. duryi respectively on growth and egg laying capacity of B. camerunensis and on the hatchability of B. camerunensis egg masses was compared. No differences were found besides an indication of a growth promoting effect of B. camerunensis conditioning. B. camerunensis egg masses were found to be much more sensitive to mechanical interference from snails than were H. duryi egg masses. Both juvenile and adult H. duryi caused a great reduction in the number of B. camerunensis egg masses and adult B. camerunensis showed a great tendency to destroy its own egg masses. Newly hatched B. camerunensis appeared to be destroyed to a greater extent by adult H. duryi than by adult B. camerunensis.     

Gene regulation of shell banding in a land snail from Israel

The Israeli land snail, Xeropicta vestalis, offers a particularly clear example of gene regulation in relation to natural selection, in that within each population the appropriate phenotype is generated only at the correct part of the animals life cycle, and a contrasting phenotype develops when the forces of natural selection change. In the mountains of Jerusalem, where the winter is cold, the shells are dark. Westwards, towards the coastal plain where the winter is warmer, the shells gradually become paler. As dark shells absorb more radiation than pale ones, this clinal variation in morph frequencies can be explained in thermal terms. (Banded shells are also more cryptic than non-banded shells, so that in the mountains visual selection by predators may be an additional force which favours dark shells.) Xeropicta vestalis is an annual, semelparous species: the snails hatch in winter, become mature within one year, reproduce, and then die. In the coastal plain the snails are active throughout most of the year, and they have a long period in which to grow to reach adult size. In the mountains and hills, on the other hand, the snails are active for only a very short period. They spend most of their lives as small snails, in a state of aestivation. Xeropicta vestalis must be dark in mountains because when it finally awakens, it must very rapidly and hastily reach reproduction size. A dark shell, by speeding up temperature-dependent processes in this critical stage, assists the snail to mature rapidly. Shell darkness varies with age: in the mountains and hills the shells are moderately dark when they hatch, but become darker whilst growing in early winter. In the coastal plain also, the snails are moderately dark when they hatch; but here they become paler, whilst growing in winter and spring. In both cases, each snail is darker in the colder months and paler in the hot ones. A strategy of gene regulation of shell colour is thus favoured when the subsequent forces of the environment are very contrasting in their direction, very severe—yet also very predictable.     

Inheritance of the background shell color in the snails Littorina obtusata (Gastropoda,Littorinidae)

We investigated in a gastropod mollusk Littorina obtusata (L. obtusata) the inheritance of back-ground shell coloration of the shell, which arises on the basis of three pigments: purple, orange, and yellow. We found that the hypothesis on polyallelic inheritance, as in the genus Cepaea, cannot explain the inheritance of shell color in periwinkles. We demonstrated that a separate genetic system is responsible for incorporation of each pigment into the shell. The composition of these genetic systems includes at least two genes each in the case of the yellow and purple pigments. Our analysis shows that caution should be applied when extending the results obtained in the studies of the Cepaea genus to the other species of gastropods.     

Température et plasticité du chromatisme de la coquille chez le mollusque pulmoné Helix aspersa

Phenotypic plasticity of shell pigmentation in the land snail Helix aspersa was investigated with the help of image processing followed by pixel counting for each colour present on the shell. Thus, the use of three different growth temperatures under artificial conditions showed, for the two subspecies considered, a continuous colour gradient from red to dark of the shell banding. Darker pigmentation characterized low temperatures while red bands were related to the higher temperature (25 °C). No interpretable difference appeared between the two forms.     

The combined effect of isolation and Fasciola hepatica infection on the life history traits of Fossaria cubensis.

Life history traits of Fossaria cubensis were compared between isolated and paired snails after infection with three miracidia of Fasciola hepatica. Four experimental groups were tested: isolated-unexposed, paired-unexposed, isolated-infected, and paired-infected. A repeated-measures ANOVA showed statistically significant interactions among isolation, infection, and age effects for shell size, number of egg masses per snail, number of eggs per snail, and number of viable eggs per snail. Isolated-unexposed snails exhibited the higher values of these variables and those of survival and finite and intrinsic rates of natural increase. Infection stimulated shell growth during the prepatent period, but differences were present only in paired snails since isolation causes a similar effect. Reproduction, in terms of the number of egg masses per snail and the number of eggs per mass per snail, decreases in the presence of parasitic infection, whereas isolation stimulates it. These effects were observed from early stages of infection.     

Silent assassins: predation of native New Zealand trichopteran eggs by non-native freshwater gastropods

Aquatic insects that oviposit on rocks may evolve strategies to mitigate loss of eggs to opportunistic predation by common native grazers, but such strategies may be ineffective against non-native grazers. We tested whether the eggs of common New Zealand caddisfly families (Hydrobiosidae and Hydropsychidae) that oviposit on rocks were more susceptible to predation by native or non-native snails. The native snail Potamopyrgus antipodarum (Grey, 1843) and non-native snails Physa acuta Draparnaud, 1805 and Pseudosuccinea columella (Say, 1817) were presented with caddisfly egg masses and the number of individual eggs consumed was recorded after 24 and 48 hours. The larger Pseudosuccinea readily consumed the eggs, especially spumaline-encapsulated Hydrobiosidae eggs. Physa consumed very few Hydrobiosidae eggs and no Hydropsychidae eggs, whereas Potamopyrgus did not consume eggs of either family. The trichopteran egg masses tested did not succumb to predation by the native Potamopyrgus but are highly vulnerable to predation by a larger non-native snail.     

A laboratory assessment of the potential molluscicidal activity of some Nigerian plant species used as anthelmintics

Laboratory studies were conducted to determine the potential molluscicidal activity of five Nigerian plant species used in alternative medicine practice as anthelmintics. Laboratory-bred adult Biomphalaria pfeifferi, the snail host of Schistosoma mansoni in Nigeria, and their viable 0- to 24-hour-old egg masses were separately exposed for 24 hours, in replicated bioassays, to 7.81–4 000 mg l^?1 concentrations of crude ethanolic extracts of the leaves and roots of Annona senegalensis, Anogeissus leiocarpus, Crotalaria retusa, Dalbergia sissoo and Vernonia amygdalina. Fruits and stem bark of D. sissoo were also investigated. The 24-hour LC50 and LC90 values of extracts for target organisms were calculated using probit analysis. Susceptibility of target organisms to extracts varied with the plant species and morphological part. Egg masses and adult snails were most susceptible to D. sissoo fruit extract (LC90 values of 89.29 and 74.33 mg l^?1, respectively) and least susceptible to V. amygdalina leaf extract (LC90 values of 11 864.09 and 6 241.61 mg l^?1, respectively). Concentration-dependent behavioural responses and embryo deaths at the gastrula/exogastrula and/or prehatch snail stages of development were observed in exposed snails and egg masses, respectively. Tested extracts of A. senegalensis, A. leiocarpus, C. retusa and V. amygdalina did not show appreciable toxicities to B. pfeifferi egg masses or adults. For demonstrating significant molluscicidal activity (LC90 < 100 mg l^?1) and toxicities to egg masses, we recommend further studies on the ethanolic extracts of D. sissoo fruits and roots.     

Quantitative measures and 3D shell models reveal interactions between bands and their position on growing snail shells

The nature of shell growth in gastropods is useful because it preserves the ontogeny of shape, colour, and banding patterns, making them an ideal system for understanding how inherited variation develops, is established and maintained within a population. However, qualitative scoring of inherited shell characters means there is a lack of knowledge regarding the mechanisms that control fine variation. Here, we combine empirical measures of quantitative variation and 3D modeling of shells to understand how bands are placed and interact. By comparing five‐banded Cepaea individuals to shells lacking individual bands, we show that individual band absence has minor but significant impacts upon the position of remaining bands, implying that the locus controlling band presence/absence mainly acts after position is established. Then, we show that the shell grows at a similar rate, except for the region below the lowermost band. This demonstrates that wider bands of Cepaea are not an artifact of greater shell growth on the lower shell; they begin wider and grow at the same rate as other bands. Finally, we show that 3D models of shell shape and banding pattern, inferred from 2D photos using ShellShaper software, are congruent with empirical measures. This work therefore establishes a method that may be used for comparative studies of quantitative banding variation in snail shells, extraction of growth parameters, and morphometrics. In the future, studies that link the banding phenotype to the network of shell matrix proteins involved in biomineralization and patterning may ultimately aid in understanding the diversity of shell forms found in molluscs.     

Y-linked Mendelian inheritance of giant and dwarf male morphs in shell-brooding cichlids

Behavioural variation among conspecifics is typically contingent on individual state or environmental conditions. Sex-specific genetic polymorphisms are enigmatic because they lack conditionality, and genes causing adaptive trait variation in one sex may reduce Darwinian fitness in the other. One way to avoid such genetic antagonism is to control sex-specific traits by inheritance via sex chromosomes. Here, controlled laboratory crossings suggest that in snail-brooding cichlid fish a single locus, two-allele polymorphism located on a sex-linked chromosome of heterogametic males generates an extreme reproductive dimorphism. Both natural and sexual selection are responsible for exceptionally large body size of bourgeois males, creating a niche for a miniature male phenotype to evolve. This extreme intrasexual dimorphism results from selection on opposite size thresholds caused by a single ecological factor, empty snail shells used as breeding substrate. Paternity analyses reveal that in the field parasitic dwarf males sire the majority of offspring in direct sperm competition with large nest owners exceeding their size more than 40 times. Apparently, use of empty snail shells as breeding substrate and single locus sex-linked inheritance of growth are the major ecological and genetic mechanisms responsible for the extreme intrasexual diversity observed in Lamprologus callipterus.     

QUANTITATIVE GENETICS OF SHELL FORM OF AN INTERTIDAL SNAIL: CONSTRAINTS ON SHORT-TERM RESPONSE TO SELECTION

We investigated the genetic and environmental determinants of shell form in an intertidal snail (Prosobranchia: Littorina sp.) to identify constraints on the short-term response to selection. Our quantitative genetic parameters were estimated from a half-sib experimental design using 288 broods of snails. Each brood was divided into two treatments differing in snail population density, and therefore in grazing area per snail. Differences in population density induced marked differences in shell form. Snails in the low density treatment grew faster and had lighter shells with narrower whorls and narrower apertures than their siblings at high density. Despite this environmental plasticity in shell shape we found significant additive genetic variance for components of shell shape. We discuss two mechanisms that may maintain additive genetic variance for shell shape in intertidal snail populations: migration between environments with different selective pressures and migration between environments with different mean growth rates. We also estimated a genetic variance-covariance matrix for shell form traits and used the matrix to identify constraints on the short-term response to selection. We predict the rate of response to selection for predator-resistant morphology such as would occur upon invasion of predatory crabs. The large negative genetic correlation between relative spire height and shell weight would facilitate simultaneous selection for a lower spire and a heavier shell, both of which would increase resistance to predatory crabs.     

Report of a cohesive gelatinous egg mass produced by a tropical marine bivalve

Abstract. Gelatinous egg masses are common in a number of animal phyla. However, they are virtually unknown in marine bivalves, with structures that could be thought of as gelatinous egg masses being reported for only five species. We describe the gelatinous egg mass and intracapsular development in the tropical lucinid Phacoides pectinatus. The embryos developed within individual capsules embedded in a large flimsy, spherical mass. Swimming veligers hatch at 198 μm shell length. They did not feed, settled within several days of hatching, and metamorphosis was completed within 2 weeks of hatching. Gelatinous egg masses might be detected in members of more lucinid species if studies of development included field or in vivo observations of reproduction in addition to producing embryos by stripping the gonads.     

Inheritance of seed coat color genes in <Emphasis Type="Italic">Brassica napus</Emphasis> (L.) and tagging the genes using SRAP,SCAR and SNP molecular markers

Seed coat color inheritance in Brassica napus was studied in F₁, F₂, F₃ and backcross progenies from crosses of five black seeded varieties/lines to three pure breeding yellow seeded lines. Maternal inheritance was observed for seed coat color in B. napus, but a pollen effect was also found when yellow seeded lines were used as the female parent. Seed coat color segregated from black to dark brown, light brown, dark yellow, light yellow, and yellow. Seed coat color was found to be controlled by three genes, the first two genes were responsible for black/brown seed coat color and the third gene was responsible for dark/light yellow seed coat color in B. napus. All three seed coat color alleles were dominant over yellow color alleles at all three loci. Sequence related amplified polymorphism (SRAP) was used for the development of molecular markers co-segregating with the seed coat color genes. A SRAP marker (SA12BG18388) tightly linked to one of the black/brown seed coat color genes was identified in the F₂ and backcross populations. This marker was found to be anchored on linkage group A9/N9 of the A-genome of B. napus. This SRAP marker was converted into sequence-characterized amplification region (SCAR) markers using chromosome-walking technology. A second SRAP marker (SA7BG29245), very close to another black/brown seed coat color gene, was identified from a high density genetic map developed in our laboratory using primer walking from an anchoring marker. The marker was located on linkage group C3/N13 of the C-genome of B. napus. This marker also co-segregated with the black/brown seed coat color gene in B. rapa. Based on the sequence information of the flanking sequences, 24 single nucleotide polymorphisms (SNPs) were identified between the yellow seeded and black/brown seeded lines. SNP detection and genotyping clearly differentiated the black/brown seeded plants from dark/light/yellow-seeded plants and also differentiated between homozygous (Y2Y2) and heterozygous (Y2y2) black/brown seeded plants. A total of 768 SRAP primer pair combinations were screened in dark/light yellow seed coat color plants and a close marker (DC1GA27197) linked to the dark/light yellow seed coat color gene was developed. These three markers linked to the three different yellow seed coat color genes in B. napus can be used to screen for yellow seeded lines in canola/rapeseed breeding programs.     

Endozoic algae in shelled gastropods — a new symbiotic association in coral reefs?

Live algae were found in the hepatopancreas and gonads of the Red Sea snail Strombus tricornis. These organs are constantly concealed within the upper whorls of the snail's shell. Light penetration was 5–15% of the incident light reaching the shell. Pigment analysis indicated the presence of chlorophyll a, c and peridinin, a composition resembling the Dinoflagellata. Chlorophyll a concentration in the algae was 1.18±0.36 pg chl/cell. ¹⁴C assimilation of isolated algae incubated in the light exceeded that of dark controls, demonstrating the photosynthetic activity of the endozoic algae.