Larval ecology and morphology in fossil gastropods

The shell of marine gastropods conserves and reflects early ontogeny, including embryonic and larval stages, to a high degree when compared with other marine invertebrates. Planktotrophic larval development is indicated by a small embryonic shell (size is also related to systematic placement) with little yolk followed by a multiwhorled shell formed by a free‐swimming veliger larva. Basal gastropod clades (e.g. Vetigastropoda) lack planktotrophic larval development. The great majority of Late Palaeozoic and Mesozoic ‘derived’ marine gastropods (Neritimorpha, Caenogastropoda and Heterobranchia) with known protoconch had planktotrophic larval development. Dimensions of internal moulds of protoconchs suggest that planktotrophic larval development was largely absent in the Cambrian and evolved at the Cambrian–Ordovician transition, mainly due to increasing benthic predation. The evolution of planktotrophic larval development offered advantages and opportunities such as more effective dispersal, enhanced gene flow between populations and prevention of inbreeding. Early gastropod larval shells were openly coiled and weakly sculptured. During the Mid‐ and Late Palaeozoic, modern tightly coiled larval shells (commonly with strong sculpture) evolved due to increasing predation pressure in the plankton. The presence of numerous Late Palaeozoic and Triassic gastropod species with planktotrophic larval development suggests sufficient primary production although direct evidence for phytoplankton is scarce in this period. Contrary to previous suggestions, it seems unlikely that the end‐Permian mass extinction selected against species with planktotrophic larval development. The molluscan classes with highest species diversity (Gastropoda and Bivalvia) are those which may have planktotrophic larval development. Extremely high diversity in such groups as Caenogastropoda or eulamellibranch bivalves is the result of high phylogenetic activity and is associated with the presence of planktotrophic veliger larvae in many members of these groups, although causality has not been shown yet. A new gastropod species and genus, Anachronistella peterwagneri, is described from the Late Triassic Cassian Formation; it is the first known Triassic gastropod with an openly coiled larval shell.     

Potamopyrgus antipodarum(Mollusca:Hydrobiidae) in continental aquatic gastropod communities: impact of salinity and trematode parasitism

The structure of gastropod communities was examined from January to June 1999 in four sites of the streams of Mont Saint-Michel Bay along a gradient of salinity, and the occurrence of larval trematodes infecting snails was studied. Abundance and species richness of gastropods increased from polyhaline (95 snails, 1 species) to oligohaline waters (6672 snails, 6 species). Whatever the salinity, the most abundant species was Potamopyrgus antipodarum, an invasive non-indigenous species that represented 80% of the gastropods. Only one male was found in P. antipodarum populations suggesting a predominantly parthenogenetic mode of reproduction. Among 7218 gastropods collected, 1.2% were infected by larval trematodes: 5 species in Lymnaea peregra (4.4%), 4 species in Planorbis planorbis (12.0%), one echinostome in Physa acuta (0.2%), and a new species of Sanguinicola in P. antipodarum (0.5%). This is the first record of infected P. antipodarum in Europe. No parasites were found in polyhaline waters. The prevalence per host population varied from 0 to 100% depending on time of collection, salinity and host species. In the lowest-salinity site, abundance of gastropods and prevalence of trematodes were negatively correlated. The dominance of P. antipodarum in the gastropod communities is discussed in relation with euryhalinity, parthenogenesis and weak rate of parasitism.     

Mining for SSRs and FDMs from expressed sequence tags of Camellia sinensis

Simple Sequence Repeats (SSRs) developed from Expressed Sequence Tags (ESTs), known as EST-SSRs are most widely used and potentially valuable source of gene based markers for their high levels of crosstaxon portability, rapid and less expensive development. The EST sequence information in the publicly available databases is increasing in a faster rate. The emerging computational approach provides a better alternative process of development of SSR markers from the ESTs than the conventional methods. In the present study, 12,851 EST sequences of Camellia sinensis, downloaded from National Center for Biotechnology Information (NCBI) were mined for the development of Microsatellites. 6148 (4779 singletons and 1369 contigs) non redundant EST sequences were found after preprocessing and assembly of these sequences using various computational tools. Out of total 3822.68 kb sequence examined, 1636 (26.61%) EST sequences containing 2371 SSRs were detected with a density of 1 SSR/1.61 kb leading to development of 245 primer pairs. These mined EST-SSR markers will help further in the study of variability, mapping, evolutionary relationship in Camellia sinensis. In addition, these developed SSRs can also be applied for various studies across species.     

Crepidula fornicata is not a first intermediate host for trematodes: who is?

Trematode larvae must generally invade a molluscan intermediate host, usually a gastropod, before they can reach reproductive maturity in another definitive host. The research literature to date has focused almost exclusively on the documented specificity between particular trematode species and particular molluscan hosts; little attention has been paid to gastropod species that do not appear to serve as hosts. We sampled Rhode Island and Massachusetts populations of the marine gastropod Crepidula fornicata to determine whether this widespread species serves as a first intermediate host for trematodes. We also sampled from the same habitat populations of Littorina littorea and Ilyanassa obsoleta, gastropods known to serve as first intermediate hosts for several trematode species. All individuals were examined by dissection for the presence of sporocysts, rediae, or developing cercariae. Although 4-28% of L. littorea (N=112) and I. obsoleta (N=84) were infected by larvae of at least one trematode species, no individuals of C. fornicata sampled from the same locations were so infected (N=136). A survey of the Biological Abstracts computer database indicates that snails in only about 10% of marine gastropod families are known to serve as first intermediate hosts for trematodes. We suggest that more attention be paid to marine gastropods that appear not to be infected by trematode miracidia. Such species may productively serve as new models for understanding trematode host specificity and gastropod resistance to infection.     

Factors affecting the structure of epiphytic gastropod communities in the St. Lawrence River (Quebec,Canada)

A study of epiphytic gastropods associated with two submerged macrophytes (Myriophyllum spicatum and Vallisneria americana) was conducted at a soft and at a hard water site in the St. Lawrence River during two non-consecutive years in order to compare effects of macrophyte species and biomass, site and year in gastropod community structure. There was no effect of macrophyte species on total gastropod abundance nor on gastropod diversity, and few gastropod species showed a marked preference for either macrophyte species. Inter-site and inter-annual variations in diversity, total gastropod abundance and gastropod community structure were greater than variations among macrophytes. However, analysis of variations of epiphytic gastropod communities in one of the two sites shows that abiotic factors are important in explaining epiphytic gastropod distribution. Our results and results of other studies on gastropod population dynamics in the St. Lawrence River indicate that intra- and interspecific competition between gastropods is important on macrophytes and that they must be carefully considered in order to understand epiphytic community structure and population dynamics.     

Molluscs in the marine curio and souvenir trade in NE Brazil: species composition and implications for their conservation and management

Marine molluscs are sold all over the world as curios and souvenirs. This market encompasses about 5,000 species of molluscs (bivalves and gastropods) worldwide, but the mollusc species involved in this trade are undocumented in all but a very small number of countries. The present study provides a baseline assessment of the mollusc species used in the manufacture of crafts, marine curiosities and souvenirs in northeastern Brazil. A total of 126 species (41 bivalves and 85 gastropods) were found to be sold individually as decorative pieces or incorporated into utilitarian objects. Overall, the families most represented were Veneridae bivalves (9 species) and the gastropods Conidae and Strombidae (8 species each). Large gastropod shells were observed in all of the localities surveyed (mainly as part of table lamps). The main species used were the King helmet Cassis tuberosa, the Hebrew volute Voluta ebraea, and the Goliath conch Eustrombus goliath. The majority of the species involved are harvested from the Atlantic Ocean (68%), but many are imported from Indo-Pacific countries, such as cowries Monetaria moneta. Among the species harvested in the Atlantic Ocean, 11.2% are endemic to Brazil as, for example, the Brazilian chank (Turbinella laevigata). The majority of the species sold in northeastern Brazil tend to inhabit shallow habitats, which facilitates their capture. As there are no official statistics available concerning the marine curio and souvenir trade in Brazil, some species may be endangered. Harvesting regulations should include minimum capture sizes, capture quotas, specific periods for harvesting, and the use of non-destructive techniques.     

High bioaccumulation of cadmium and other metals in Patagonian edible gastropods

High concentrations of metals are accumulated by edible marine resources, affecting human health. Marine gastropods have been commercially captured and consumed in Argentina for decades without official regulations or studies to guarantee their safety. In this study, metals (cadmium, aluminium, iron, zinc, copper and lead) were analysed for the first time in the edible marine gastropods Buccinanops globulosus, Adelomelon ancilla and Trophon geversianus and their surrounding sediments. Taking into account the maximum levels recommended by national and international regulations for food safety, we recommend consuming only the foot of these gastropod species. We recommend not consuming these gastropods from harbour areas due to concentrations of lead in sediments and tissues. Though the target hazard quotient (THQ) of cadmium was not high for the local community, it could be higher for other populations who consume molluscs more frequently, indicating a potential health risk of chronic exposure. Monitoring of these and other contaminants should be performed in order to ensure food safety of these marine resources.     

Mesoherbivores reduce net growth and induce chemical resistance in natural seaweed populations

Herbivory on marine macroalgae (seaweeds) in temperate areas is often dominated by relatively small gastropods and crustaceans (mesoherbivores). The effects of these herbivores on the performance of adult seaweeds have so far been almost exclusively investigated under artificial laboratory conditions. Furthermore, several recent laboratory studies with mesoherbivores indicate that inducible chemical resistance may be as common in seaweeds as in vascular plants. However, in order to further explore and test the possible ecological significance of induced chemical resistance in temperate seaweeds, data are needed that address this issue in natural populations. We investigated the effect of grazing by littorinid herbivorous snails (Littorina spp.) on the individual net growth of the brown seaweed Ascophyllum nodosum in natural field populations. Furthermore, the capacity for induced resistance in the seaweeds was assessed by removing herbivores and assaying for relaxation of defences. We found that ambient densities of gastropod herbivores significantly reduced net growth by 45% in natural field populations of A. nodosum. Seaweeds previously exposed to grazing in the field were less consumed by gastropod herbivores in feeding bioassays. Furthermore, the concentration of phlorotannins (polyphenolics), which have been shown to deter gastropod herbivores, was higher in the seaweeds that were exposed to gastropod herbivores in the field. This field study corroborates earlier laboratory experiments and demonstrates that it is important to make sure that the lack of experimental field data on marine mesoherbivory does not lead to rash conclusions about the lack of significant effects of these herbivores on seaweed performance. The results strongly suggest that gastropods exert a significant selection pressure on the evolution of defensive traits in the seaweeds, and that brown seaweeds can respond to attacks by natural densities of these herbivores through increased chemical resistance to further grazing.     

Rare, threatened and alien species in the gastropod communities in the clay pit ponds in relation to the environmental factors (The Ciechanowska Upland, Central Poland)

The objectives of the present survey were to formulate a zoocenological analysis of the gastropod communities in clay pit ponds, as well as to determine the relationships between the gastropods and their environments, and to evaluate the water habitats in terms of their ecological-conservation value. The physical and chemical parameters of water, gastropod species and macrophytes occurring in the ponds indicate mesotrophic conditions. Principal Component Analysis (PCA) reveals a correlation between gastropod density and nitrates, a correlation between gastropod density and alkalinity, and additionally shows a correlation between gastropod density and chlorides. The pond surface area, nitrates, chlorides concentrate, alkalinity, and substratum have all affected these gastropod communities. The occurrence of the first permanent population of Ferrissia clessiniana (Jickeli, 1882) was recorded in Poland. Valvata naticina Menke 1845 has become a critically rare species (CR) due to the pollution of water environments. Five other species, e.g. Anisus vorticulus (Troschel, 1834) and Planorbis carinatus O.F. Müller, 1774 are recorded on the Polish Red List of Species. The clay pit ponds of the Ciechanowska Upland, because of their distinctive environmental features, provide a refuge where a number of rare, threatened and alien gastropod species live.     

Clarifying marine invasions with molecular markers: an illustration based on mtDNA from mistaken calyptraeid gastropod identifications

Species invasions are occurring at an increasing rate in coastal environments. Accurately identifying introductions is a critical issue to take full advantage of the new invasion databases. Further, life history differences between morphologically comparable species may require that different management strategies be instigated to effectively control different species. Facing this problem, we used molecular approaches and documented a case of mistaken identification in a group of marine invertebrates, the calyptraeid gastropods. Members of this group have repeatedly and successfully invaded new habitats after anthropogenic introduction, especially in estuaries and bays on the west coast of the United States of America. For example, Crepidula fornicata, native to the east coast of the USA, has been reported from at least five USA west coast estuaries. We sequenced a fragment of the COI gene of a sample of putative C. fornicata from Humboldt Bay, California. By constructing a phylogeny of these and other calpytraeid gastropod sequences, we discovered that the individuals were C. convexa, the convex slippershell. In contrast to C. fornicata, C. convexa has large, demersal eggs and larvae are well developed at hatching. Its potential for dispersal is therefore lower as compared to C. fornicata and therefore any strategy to manage the invasion should take this into account. In the present study, we demonstrated the utility of molecular tools that can be used by non-taxonomic experts, to quickly and accurately identify alien species – an important first step in any study of invasion biology.     

Development of chickpea EST-SSR markers and analysis of allelic variation across related species

Despite chickpea being the third important grain legume, there is a limited availability of genomic resources, especially of the expressed sequence tag (EST)-based markers. In this study, we generated 822 chickpea ESTs from immature seeds as well as exploited 1,309 ESTs from the chickpea database, thus utilizing a total of 2,131 EST sequences for development of functional EST-SSR markers. Two hundred and forty-six simple sequence repeat (SSR) motifs were identified from which 183 primer pairs were designed and 60 validated as functional markers. Genetic diversity analysis across 30 chickpea accessions revealed ten markers to be polymorphic producing a total of 29 alleles and an observed heterozygosity average of 0.16 thereby exhibiting low levels of intra-specific polymorphism. However, the markers exhibited high cross-species transferability ranging from 68.3 to 96.6% across the six annual Cicer species and from 29.4 to 61.7% across the seven legume genera. Sequence analysis of size variant amplicons from various species revealed that size polymorphism was due to multiple events such as copy number variation, point mutations and insertions/deletions in the microsatellite repeat as well as in the flanking regions. Interestingly, a wide prevalence of crossability-group-specific sequence variations were observed among Cicer species that were phylogenetically informative. The neighbor joining dendrogram clearly separated the chickpea cultivars from the wild Cicer and validated the proximity of C. judaicum with C. pinnatifidum. Hence, this study for the first time provides an insight into the distribution of SSRs in the chickpea transcribed regions and also demonstrates the development and utilization of genic-SSRs. In addition to proving their suitability for genetic diversity analysis, their high rates of transferability also proved their potential for comparative genomic studies and for following gene introgressions and evolution in wild species, which constitute the valuable secondary genepool in chickpea. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Towards an efficient computational mining approach to identify EST-SSR markers

Microsatellites are the markers of choice due to their high abundance reproducibility, degree of polymorphism and co-dominant nature. These are mainly used for studying the genetic variability in different species and Marker assisted selection. Expressed Sequence Tags (ESTs) serve as the main resource for Simple Sequence Repeats (SSRs). The computational approach for detecting SSRs and developing SSR markers from EST-SSRs is preferred over the conventional methods as it reduces time and cost to a great extent. The available EST sequence databases, various web interfaces and standalone tools provide the platform for an easy analysis of the EST sequences leading to the development of potential EST-SSR Markers. This paper is an overview of in silico approach to develop SSR Markers from the EST sequence using some of the most efficient tools that are available freely for academic purpose.     

PROTEINS INVOLVED IN MEMBRANE TRANSPORT BETWEEN THE ER AND THE GOLGI APPARATUS: 21 PUTATIVE PLANT HOMOLOGUES REVEALED BY DBEST SEARCHING

Numerous proteins have been identified in yeast and mammalian cells which are involved in trafficking between the endoplasmic reticulum and the Golgi apparatus. A great number of partial cDNA sequences now available from the two major plant model species, Arabidopsis thaliana and Oryza sativa, makes it possible to identify putative plant homologues of known genes/proteins from non-plant species. The authors used this approach to screen the database of Expressed Sequence Tags (dbEST) in order to detect plant homologues of proteins involved in membrane transport between ER and Golgi. Availability of these partial sequences will facilitate the screening of cDNA and genomic libraries otherwise performed using heterologous probes derived from animal and yeast genes. As the plant Golgi complex differs in many respects from its mammalian and yeast counterparts, the dbEST clones found can be directly used for various functional assays (immunoprecipitation, two-hybrid analysis, transgenic plants etc.) to test the exact roles of the encoded proteins and identify their functional partners, some of which may be specific for plants.     

SSR mining in coffee tree EST databases: potential use of EST-SSRs as markers for the Coffea genus

Expressed sequence tags (ESTs) from Coffea canephora leaves and fruits were used to search for types and frequencies of simple sequence repeats (EST–SSRs) with a motif length of 1–6 bp. From a non-redundant (NR) EST set of 5,534 potential unigenes, 6.8% SSR-containing sequences were identified, with an average density of one SSR every 7.73 kb of EST sequences. Trinucleotide repeats were found to be the most abundant (34.34%), followed by di- (25.75%) and hexa-nucleotide (22.04%) motifs. The development of unique genic SSR markers was optimized by a computational approach which allowed us to eliminate redundancy in the original EST set and also to test the specificity of each pair of designed primers. Twenty-five EST–SSRs were developed and used to evaluate cross-species transferability in the Coffea genus. The orthology was supported by the amplicon sequence similarity and the amplification patterns. The >94% identity of flanking sequences revealed high sequence conservation across the Coffea genus. A high level of polymorphic loci was obtained regardless of the species considered (from 75% for C. liberica to 86% for C. canephora). Moreover, the polymorphism revealed by EST–SSR was similar to that exposed by genomic SSR. It is concluded that Coffea ESTs are a valuable resource for microsatellite mining. EST-SSR markers developed from C. canephora sequences can be easily transferred to other Coffea species for which very little molecular information is available. They constitute a set of conserved orthologous markers, which would be ideal for assessing genetic diversity in coffee trees as well as for cross-referencing transcribed sequences in comparative genomics studies.     

Prasiola (Prasiolales,Trebouxiophyceae) in Japan: a survey of freshwater populations and new records of marine taxa

Recent collections from marine and freshwater locations have enabled the investigation of diversity of Prasiola in Japan. Sequence data from the rbc L and tuf A markers revealed the presence of three marine species and one freshwater species. Prasiola delicata was confirmed to occur on Daikokujima, Prasiola calophylla was found for the first time in Japan from Hokkaido, and a species within the P. meridionalis/linearis/stipitata complex was found on both Hokkaido and Daikokujima. Collections from a range of populations of freshwater Prasiola, identified here as P. japonica, were found to be conspecific and identical in rbc L and tuf A sequences to freshwater collections from Nepal, Korea, and China.     

Long-term dynamics and community structure of freshwater gastropods exposed to parasitism and other environmental stressors

1. Freshwater communities are subject to various abrupt environmental disturbances and increasing pollution levels in their habitat. According to the nature and the periodicity of disturbances (i.e. punctuated, recurrent or continuous), one may expect different kinds of response of the target gastropod communities. 2. The gastropod community of a French lake has been investigated for 10 years to study its responses to three main environmental disturbances: supra‐seasonal droughts, recurrent proliferations of cyanobacteria and parasitism by trematodes. The relationship between this latter factor, considered as a continuous stressor, and the gastropod community was a particular focus. 3. A total of 13 280 gastropods belonging to 17 species (mostly Pulmonata with Planorbidae as the dominant family) have been sampled in the lake over the 10‐year time series. Species composition of the gastropod community varied strongly during the study period as did species richness (0–14, mean of 6.5 ± 0.4 species) and abundance of gastropods (0–4456, mean of 192 ± 72 individuals). Trematode larvae belonging to 11 morphotypes of cercariae were detected in 15 of the 17 species of gastropods and had a total prevalence of 2.9%. 4. Droughts (punctuated stressor) were responsible for the disappearance of nearly all gastropod species, and thus represent a severe environmental stressor for them. Nevertheless, the extent of this stressor was limited since the gastropod community was highly resilient. 5. Recurrent proliferations of toxic cyanobacteria (recurrent stressor) coincided with the large decline of the gastropod community, corresponding to a strong decrease in abundance and species richness. Intoxication of gastropods by toxic cyanobacteria was demonstrated by the bioaccumulation of microcystins in all gastropod tissues. 6. Trematode parasitism (continuous stressor) did not affect the gastropod community despite the possibly strong impact of parasites on some host species (those with monthly prevalence as high as 100% at some times). Indeed, abundance and species richness of gastropods were scarcely influenced by trematode prevalence or species richness.     

Spiders and harvestmen as gastropod predators

1. Reports are reviewed of gastropod feeding (malacophagy) by spiders and harvestmen. Although the standard textbooks on arachnids recognise the importance of gastropods as prey of harvestmen, none apparently refers to malacophagy by spiders. A review of several hundred papers on spider feeding habits revealed that species from several families kill and devour slugs and snails in the laboratory and/or field. 2. Malacophagy has been reported most frequently among mygalomorph spiders, and can make up a substantial proportion of the diets of some species, however gastropods make up an insignificant percentage of the prey of most araneomorph spiders. The spiders that eat gastropods are species with broad diets composed predominantly of arthropod prey. No species of spider appears to feed exclusively on gastropod prey. 3. Harvestmen from several families have broad diets that often include gastropods. Several species of the family Trogulidae and at least one species of the family Ischyropsalididae [Ischyropsalis hellwigi (Panzer)] are specialised gastropod predators. The trogulids are slender animals that attack the snail through the shell aperture (shell intruders). Ischyropsalis hellwigi, on the other hand, can crush snail shells with its powerful chelicerae (shell breakers). 4. The review highlights apparent convergent evolution by harvestmen and Carabidae of two mutually exclusive morphologies found among gastropod predators. It also suggests that there is an urgent need for systematic studies to be conducted to establish the extent and ecological importance of malacophagy in natural and anthropogenically altered habitats.