Laboratory spawning of the purple snail Plicopurpura pansa (Gastropoda: Muricidae)

The spawning of the muricid gastropod Plicopurpura pansa in the laboratory at 22-23 degrees C is described. Females deposited 1-20 capsules daily for at least 20 weeks, and produced up to 150 capsules each per spawning season. During spawning, egg clusters were formed consisting of hundreds of capsules of different ages deposited by different females. Each egg capsule contained an average of 436 embryos (+/- s.d. 213.6, range: 95-1092, n=50). Embryos developed without nurse eggs. After six to eight weeks of intracapsular, lecithotrophic development, planktotrophic veligers hatched with two fully developed velar lobes.     

Speciation and diversity on tropical rocky shores: a global phylogeny of snails of the genus Echinolittorina

A phylogenetic approach to the origin and maintenance of species diversity ideally requires the sampling of all species within a clade, confirmation that they are evolutionarily distinct entities, and knowledge of their geographical distributions. In the marine tropics such studies have mostly been of fish and reef-associated organisms, usually with high dispersal. In contrast, snails of the genus Echinolittorina (Littorinidae) are restricted to rocky shores, have a four-week pelagic development (and recorded dispersal up to 1400 km), and show different evolutionary patterns. We present a complete molecular phylogeny of Echinolittorina, derived from Bayesian analysis of sequences from nuclear 28S rRNA and mitochondrial 12S rRNA and COI genes (nodal support indicated by posterior probabilities, maximum likelihood, and neighbor-joining bootstrap). This consists of 59 evolutionarily significant units (ESUs), including all 50 known taxonomic species. The 26 ESUs found in the Indo-West Pacific region form a single clade, whereas the eastern Pacific and Atlantic species are basal. The earliest fossil occurred in the Tethys during the middle Eocene and we suggest that the Indo-West Pacific clade has been isolated since closure of the Tethyan seaway in the early Miocene. The geographical distributions of all species (based on more than 3700 locality records) appear to be circumscribed by barriers of low temperature, unsuitable sedimentary habitat, stretches of open water exceeding about 1400 km, and differences in oceanographic conditions on the continuum between oceanic and continental. The geographical ranges of sister species show little or no overlap, indicating that the speciation mode is predominantly allopatric. Furthermore, range expansion following speciation appears to have been limited, because a high degree of allopatry is maintained through three to five branching points of the phylogeny. This may be explained by infrequent long-distance colonization, habitat specialization on the oceanic/continental gradient, and perhaps by interspecific competition. In the eastern Pacific plus Atlantic we identify five cases of divergence on either side of the Isthmus of Panama, but our estimates of their ages pre-date the emergence of the Isthmus. There are three examples of sister relationships between species in the western Atlantic and eastern Atlantic, all resulting from dispersal to the east. Within the Indo-West Pacific, we find no geographical pattern of speciation events; narrowly endemic species of recent origin are present in both peripheral and central parts of the region. Evidence from estimated divergence times of sister species, and from a plot of the number of lineages over time, suggest that there has been no acceleration of diversification during the glacio-eustatic cycles of the Plio-Pleistocene. In comparison with reefal organisms, species of Echinolittorina on rocky shores may be less susceptible to extinction or isolation during sea-level fluctuations. The species richness of Echinolittorina in the classical biogeographic provinces conforms to the common pattern of highest diversity (11 species) in the central "East Indies Triangle" of the Indo-West Pacific, with a subsidiary focus in the eastern Pacific and western Atlantic, and lowest diversity in the eastern Atlantic. The diversity focus in the East Indies Triangle is produced by a mosaic of restricted allopatric species and overlap of a few widespread ones, and is the result of habitat specialization rather than historical vicariance. This study emphasizes the plurality of biogeographic histories and speciation patterns in the marine tropics.     

Effects of foliaceous varices on the mechanical properties of Chicoreus dilectus (Gastropoda: Muricidae)

Removal of the foliaceous varices from the shells of Chicoreus dilectus had no significant effect on the crushing strength (load at failure) of the shells, but decreased the total work necessary to break the shells by 35–50%. This effect was due to the removal of varices as an architectural feature of the shell rather than simply to the removal of shell material per se ; varicate shells had a mean work to failure per unit shell mass 1–6–2–3 times greater than avaricate shells. Because the strength of gastropod shells often exceeds the maximum forces that durophagous predators may bring to bear on them, work to failure may be a more important factor in the design of gastropod shells than breaking load.     

Cytogenetics of Bolinus brandaris and phylogenetic inferences within the Muricidae (Mollusca: Gastropoda)

The purple dye murex, Bolinus brandaris (Linnaeus, 1758), is a muricid gastropod common throughout the Mediterranean and along the Moroccan and Portuguese Atlantic coasts. In the present study, we confirmed the diploid chromosome number of 2 n  = 70 for this species, and established for the first time the karyotype, which comprised 12 metacentric, 15 submetacentric and eight subtelocentric chromosome pairs. To facilitate cytotaxonomic comparisons, we carried out a comparative karyological analysis through multidimensional scaling between B. brandaris and three other 2 n  = 70 muricid species ( Hexaplex trunculus , Ocenebra erinaceus , and Stramonita haemastoma ) for which chromosomal measurements have been previously published. The interpretation of the ideograms and the statistical analysis highlighted the closest similarity of B. brandaris and H. trunculus compared to S. haemastoma and O. erinaceus . Indeed, B. brandaris and H. trunculus showed the smallest dissimilarities both for relative length and arm ratio, with O. erinaceus presenting intermediate values, whereas the highest dissimilarities were found between H. trunculus and S. haemastoma for both data. The karyotypes of B. brandaris and H. trunculus (subfamily Muricinae) presented the highest proportions of metacentric chromosomes compared to the other two muricids analysed, suggesting that those karyotypes could be considered primitive within the 2 n  = 70 Muricidae studied so far.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 96 , 185–193.     

Bridging gaps in the molecular phylogeny of the Lymnaeidae (Gastropoda: Pulmonata), vectors of Fascioliasis

Background  

Lymnaeidae snails play a prominent role in the transmission of helminths, mainly trematodes of medical and veterinary importance (e.g., Fasciola liver flukes). As this family exhibits a great diversity in shell morphology but extremely homogeneous anatomical traits, the systematics of Lymnaeidae has long been controversial. Using the most complete dataset to date, we examined phylogenetic relationships among 50 taxa of this family using a supermatrix approach (concatenation of the 16 S, ITS-1 and ITS-2 genes, representing 5054 base pairs) involving both Maximum Likelihood and Bayesian Inference.     

A cladistic phylogeny of the genus Littorina (Gastropoda): implications for evolution of reproductive strategies and for classification

Gross anatomical characters of all 18 species of Littorina are used to construct a phylogenetic hypothesis for the genus, by the method of cladistic analysis. The resulting cladogram suggests that of the four subgenera of Littorina, one (Littorina) is paraphyletic. It is uncertain whether the genus Mainwaringia should be included in Littorina. It is shown that the non-planktotrophic Littorina species in the northern Atlantic comprise a monophyletic group, with the sister-species L. kurila and/or L. subrotundata in the northern Pacific. Invasion of the Atlantic by a minimum of two Pacific species, across the Arctic migration route established during the late Cenozoic, is sufficient to account for the modern distribution of the subgenera Littorina and Neritrema. The importance of the cladogram as a basis for hypotheses of adaptation is illustrated by a discussion of spawn and development in Littorina.     

A phylogeny of the land snails (Gastropoda: Pulmonata)

We have undertaken the first large-scale molecular phylogenetic analysis of the Stylommatophora. Sequences of the ribosomal RNA gene-cluster were examined in 104 species of snails and slugs from 50 families, encompassing all the currently recognized major groups. It allows an independent test of the present classification based on morphology. At the level of families our molecular phylogeny closely supports the current taxonomy, but the deep branches within the tree do not. Surprisingly, a single assemblage including the families Achatinidae, Subulinidae and Streptaxidae lies near the base of the tree, forming a sister group to all remaining stylommatophorans. This primary division into 'achatinoid' and 'non-achatinoid' taxa is unexpected, and demands a radical reinterpretation of early stylommatophoran evolution. In particular, the Orthurethra appear to be relatively advanced within the 'non-achatinoid clade', and broadly equivalent to other super-familial clusters. This indicates that supposedly primitive features such as the orthurethran kidney are derived. The molecular tree also suggests that the origin of the Stylommatophora is much earlier than the main period of their diversification.     

The phylogeny and taxonomy of austral monodontine topshells (Mollusca: Gastropoda: Trochidae), inferred from DNA sequences

The systematics of topshells (family Trochidae) is currently unresolved: at present even the generic boundaries within this group are poorly defined. In this study, we used sequence data of two mitochondrial genes (16S and cytochrome oxidase 1, COI) and one nuclear gene (actin) to resolve the phylogeny of a closely related subgroup of the Trochidae, 30 species of largely Southern Hemisphere monodontine topshells. The phylogenies constructed revealed five well-supported generic clades: a South African clade (genus Oxystele Philippi, 1847), which lay basally to four internal Pacific clades (genera Chlorodiloma Pilsbry, 1889; Monodonta Lamarck, 1799; Austrocochlea Fischer, 1885; and Diloma Philippi, 1845). The molecular phylogenies constructed in this study shed light on previously unresolved relationships between different groups of topshells, allowing for the first time assignation (based on DNA sequence) of clearly defined, well-supported taxonomic and nomenclatural classification of monodontine topshells species. Austrocochlea crinita (Philippi, 1849), A. odontis (Wood, 1828), A. adelaidae (Philippi, 1849), and A. millelineata (Bonnet, 1864) are placed in the genus Chlorodiloma, which we resurrect from synonymy with Austrocochlea. The Japanese M. confusa Tapparone-Canefri, 1874 is treated as a separate species from M. labio (Linné, 1758). Melagraphia Gray, 1847 is synonymised with Diloma and its sole member, M. aethiops (Gmelin, 1791), along with A. concamerata (Wood, 1828), is transferred to that genus. The Juan Fernandez endemic D. crusoeana (Pilsbry, 1889) is synonymised with D. nigerrima (Gmelin, 1791). We find that morphologically cryptic species are not necessarily close genetically.     

Molecular phylogeny of the Helicodontidae and Trissexodontidae (Gastropoda)

Gómez‐Moliner, B.J., Elejalde, A.M., Arrébola, J.R., Puente, A.I., Martínez‐Ortí, A., Ruiz, A. & Madeira, MJ. (2012). Molecular phylogeny of the Helicodontidae and Trissexodontidae (Gastropoda). —Zoologica Scripta, 00, 000–000. In this study, we present a molecular phylogeny of the Trissexodontidae and Helicodontidae obtained by means of Maximum Parsimony, Neighbor Joining, Maximum Likelihood and Bayesian analyses of DNA sequences. Nearly 3 KB of sequence data of two mitochondrial genes (COI, 16S rDNA) and the nuclear rRNA gene cluster including ITS‐1, the 3′end of the 5.8S gene, the complete ITS‐2 region and 5′ end of the large subunit 28S were used to reconstruct the phylogeny of these two families. Monophyly of Trissexodontidae and Helicodontidae at the family level is well supported. A new classification of the genera in the Trissexodontidae is proposed. It includes two subfamilies: Gittenbergeriinae (monotypic for Gittenbergeria turriplana) and Trissexodontinae. The latter includes three strongly supported tribes: (i) Trissexodontini, including Mastigophallus, Trissexodon, Oestophorella and Suboestophora; (ii) Oestophorini, with Oestophora; and (iii) Caracollinini, with Caracollina, Gasulliella, Gasullia and Hatumia. The polytypic Oestophora and Suboestophora are recovered as two monophyletic genera. The anatomy of the auxiliary copulatory organs of the reproductive system is coherent with the new taxonomic interpretation of the Trissexodontidae. Further work, including some more taxa is needed to delimitate subfamilies within Helicodontidae. Finally, the addition of some sequences of other Helicoidea shows that the genus Ciliella is not closely related to Trissexodontidae, being grouped within the Hygromiidae, instead.     

Patterns of shell penetration by Chorus giganteus juveniles (Gastropoda: Muricidae) on the mussel Semimytilus algosus

Patterns are described for shell penetration by the sublittoral muricid snail Chorus giganteus during predatory attacks on the mussel Semimytilus algosus. Location, form and size of shell penetrations were observed in relation to the size of the predator. The results suggested that positions of the perforations on the mussel shells were related to size of the attacking snail. Smaller snails perforated areas near the shell ligament and in the central zone of the shell, while larger snails more frequently attacked shell borders, principally on the ventral side. These observations may be related to: (a) changes in the process of manipulation of the prey during development of the foot and the shell tooth of the predator, (b) changes in internal structure of the snails related to the shell perforation mechanism, or (c) learned behavior acquired experientially by the snails during early growth. Although in other studies of muricid penetration patterns larger boreholes made in shells of the prey were positively correlated with increasing predator size, this relation did not appear to hold with C. giganteus, as larger specimens often made relatively small shell perforations. Areas of boreholes made in the mussel shells by this snail varied from 0.01 to 1.1 mm(2), and were unusually variable in size and shape, especially when compared with literature results on bores characteristic of other muricid species.     

Egg capsules, eggs and embryos of the southwestern Atlantic gastropod Coronium coronatum (Gastropoda: Muricidae)

The egg capsules, eggs and embryos of the muricid gastropodCoronium coronatum are described for the first time. Capsulesare sessile, bulliform, semi-circular, with a plug in the dorsalcenter. Sutures split the capsule into two asymmetrical halves.Recently laid capsules showed the presence of 3639 (n = 2) uncleavednurse eggs with a diameter of 180–210 µm (mean= 197.4 ± 8.9). The number of early embryos was 9–11.The size of the embryos was 320 x 320 to 820–880 µm.Nine pre-hatching embryos of 3.94 mm (n = 8, SD = 0.32)were found inside the older capsule. SEM illustrations of embryosand radulae are provided. Comparison of shell and radula ofembryos with the protoconch and radulae of adults of C. coronatumrevealed that the capsule belongs to this species. (Received 18 March 2006; accepted 10 October 2006)     

Morphology and phylogeny of Prorocentrum caipirignum sp. nov. (Dinophyceae), a new tropical toxic benthic dinoflagellate

A new species of toxic benthic dinoflagellate is described based on laboratory cultures isolated from two locations from Brazil, Rio de Janeiro and Bahia. The morphology was studied with SEM and LM. Cells are elliptical in right thecal view and flat. They are 37–44 μm long and 29–36 μm wide. The right thecal plate has a V shaped indentation where six platelets can be identified. The thecal surface of both thecal plates is smooth and has round or kidney shaped and uniformly distributed pores except in the central area of the cell, and a line of marginal pores. Some cells present an elongated depression on the central area of the apical part of the right thecal plate. Prorocentrum caipirignum is similar to Prorocentrum lima in its morphology, but can be differentiated by the general cell shape, being elliptical while P. lima is ovoid. In the phylogenetic trees based on ITS and LSU rDNA sequences, the P. caipirignum clade appears close to the clades of P. lima and Prorocentrum hoffmannianum. The Brazilian strains of P. caipirignum formed a clade with strains from Cuba, Hainan Island and Malaysia and it is therefore likely that this new species has a broad tropical distribution. Prorocentrum caipirignum is a toxic species that produces okadaic acid and the fast acting toxin prorocentrolide.     

Relationships between height on the shore and size distributions of Thais SPP. (Gastropoda: Muricidae)

Average body sizes in a variety of marine animals have been reported to be correlated with height in the intertidal zone. This paper reports field observations on four species of Thais showing that the relationship between size distribution and intertidal height or subtidal depth is highly variable. Intertidal size gradients in Thais have been explained by a size dependent set of responses to light and gravity, but experiments reported here do not indicate that snails of different sizes behave differently with respect to the intertidal gradient per se and it is argued that, when there is a size gradient, it is produced by the direct responses of snails to factors such as shelter, temperature, desiccation, and food. It is suggested that a response to token stimuli is unlikely to evolve when the token stimuli are poorly correlated with the relevant environmental factors, especially if the latter can be monitored directly by the animal.     

A cladistic phylogeny of the family Patellidae (Mollusca: Gastropoda)

A phylogenetic hypothesis for the patellid limpets is reconstructed by cladistic analysis of morphological characters from 37 species, representing all but one of the living members of the family. Characters included in the analysis are derived from shell shape and microstructure, headfoot and pallial complex, radula and sperm. The species fall into four clades, providing the basis for a new phylogenetic classification into four monophyletic genera: Helcion (four species; southern Africa), Cymbula (eight species; southern Africa, eastern Atlantic, southern Indian Ocean), Scutellastra (17 species; southern and southwestern Africa, Australia, Indo-West Pacific, Eastern Pacific) and Patella (nine species; northeastern Atlantic and Mediterranean). The analysis suggests sister-group relationships between Helcion and Cymbula, and between Scutellastra and Patella. In combination with present-day patterns of geographical distribution, this phylogenetic hypothesis is used to discuss the historical biogeography of the Patellidae. Scutellastra may have originated in southern Africa and dispersed across the Pacific, or alternatively may be a primitively Tethyan group. Both Helcion and Cymbula appear to have originated in southern Africa, but three Cymbula species have dispersed respectively to northwest Africa, St Helena and the southern Indian Ocean. The patellids of the northeastern Atlantic form a single clade, Patella (including P. pellucida), which may have arrived by northward dispersal of an ancestor from southern Africa, or possibly by vicariance of a widespread ancestral Tethyan distribution. The known fossil record of patellids is too fragmentary to permit choice between these alternatives.     

Attack responses of the southern Australian whelk, Lepsiella vinosa (Lamarck, 1822) (Gastropoda: Muricidae), to novel bivalve prey: an experimental approach

In their native ranges, muricid gastropods feed on similar prey, often bivalves and barnacles, which they usually drill. Throughout its wide southern Australian distribution, the intertidal Lepsiella vinosa feeds on a range of prey from barnacles and littorinid gastropods in the southeast to mussels in the southwest. A number of muricids have been introduced throughout the world, either with oysters or in ship ballast water. In their new environments, they switch to native prey but their feeding responses to them have never been studied in the laboratory. The object of this study was to study the feeding of L. vinosa on a suite of non-native species. Australian Lepsiella vinosa was taken to Hong Kong, offered five different possible prey species and allowed to feed to satiation for many weeks. Replacement of consumed prey items by similar-sized conspecifics was undertaken until trends emerged. Lepsiella vinosa readily attacked thick- and thin-shelled bivalves from Hong Kong’s sandy beaches, Anomalocardia squamosa and Caecella chinensis, respectively, and from rocky shores, Septifer virgatus and Hormomya mutabilis, again respectively. It attacked them all, as it does its major prey item, Xenostrobus inconstans, in its native southwestern Australia, by either drilling or marginal probing with its proboscis. It also preferred intermediate sized prey (10–15 mm shell length), as with its natural prey. It quickly attacked the sandy shore species, and Hormomya mutabilis and Mytilopsis sallei, the latter two being closely similar in shell form and size to its natural prey X. inconstans. Hormomya mutabilis was the most favored prey, and was most similar in shell form and thickness to X. inconstans. This study therefore suggests that if introduced elsewhere, L. vinosa could radically affect intertidal community structure.     

The molluscan genus Cominella (Gastropoda: Buccinidae) at the Three Kings Islands

Abstract

Three endemic taxa of Cominella Gray are recorded from the Three Kings Islands, New Zealand. Cominella (Josepha) regalis n.sp. and C. (Josepha) quoyana griseicalx n.ssp. occur sympatrically in relatively shallow water (to c.50m), whereas C. (Eucominia) mirabilis mirabilis Powell is known only from c.180m. The history of colonisation by these taxa is hypothesised and discussed.     

Musculature of the penial complex: A new criterion in unravelling the phylogeny of Hygrophila (Gastropoda: Pulmonata)

The taxonomy of freshwater pulmonates (Hygrophila) has been in a fluid state warranting the search for new morphological criteria that may show congruence with molecular phylogenetic data. We examined the muscle arrangement in the penial complex (penis and penis sheath) of most major groups of freshwater pulmonates to explore to which extent the copulatory musculature can serve as a source of phylogenetic information for Hygrophila. The penises of Acroloxus lacustris (Acroloxidae), Radix auricularia (Lymnaeidae), and Physella acuta (Physidae) posses inner and outer layers of circular muscles and an intermediate layer of longitudinal muscles. The inner and outer muscle layers in the penis of Biomphalaria glabrata consist of circular muscles, but this species has two intermediate longitudinal layers separated by a lacunar space, which is crossed by radial and transverse fibers. The muscular wall of the penis of Planorbella duryi is composed of transverse and longitudinal fibers, with circular muscles as the outer layer. In Planorbidae, the penial musculature consists of inner and outer layers of longitudinal muscles and an intermediate layer of radial muscles. The penis sheath shows more variation in muscle patterns: its muscular wall has two layers in A. lacustris, P. acuta, and P. duryi, three layers in R. auricularia and Planorbinae and four layers in B. glabrata. To trace the evolution of the penial musculature, we mapped the muscle characters on a molecular phylogeny constructed from the concatenated 18S and mtCOI data set. The most convincing synapomorphies were found for Planorbinae (inner and outer penis layers of longitudinal muscles, three-layered wall of the penis sheath). A larger clade coinciding with Planorbidae is defined by the presence of radial muscles and two longitudinal layers in the penis. The comparative analysis of the penial musculature appears to be a promising tool in unraveling the phylogeny of Hygrophila.     

Development of indices for nonsacrificial sexing of imposex-affected Hexaplex (Trunculariopsis) Trunculus (Gastropoda: Muricidae)

The muricid Hexaplex (Trunculariopsis) trunculus lacks externalsexual dimorphism and is highly affected by imposex, which furthercomplicates its sexual identification. In this context, theaim of this study was to develop sexual indices based on thedimensions of male and imposex-affected female penis, whichcould constitute a useful, simple and nonsacrificial tool forsexing live specimens of T. trunculus. The adoption of sexualindices consisting of penis dimensions of both sexes relativeto individual size revealed a high accuracy in the sexual identificationof sacrificed specimens (>95% correct sexing). Additionally,multivariate discriminant analysis allowed correct sexing of98.8% of the original 1053 sacrificed individuals, with accuratesexual identification being higher for males (99.5%) than forimposex-affected females (97.7%). An anaesthetization experimentwas performed to investigate the effects produced by the anaesthetic(MgCl₂) on penis measurements, to test this nonsacrificial approachand to validate the previously developed sexual indices. Theanaesthetic provoked an expected enlargement in penis dimensionsbut, despite this side effect, the sexual indices developedfor sacrificed specimens were still highly successful in sexinganaesthetized T. trunculus (generally more than 95% correctsexual identification). The practical application and some limitationsof developing and employing this kind of index for the sexualidentification of T. trunculus and other imposex-affected gastropodspecies are discussed. (Received 30 May 2005; accepted 17 January 2006)     

Molecular phylogeny and systematics of Acrotoma (Gastropoda: Clausiliidae) from the Caucasus

The land snail genus Acrotoma (Clausiliidae) from the western Greater Caucasus is revised and a molecular phylogenetic analysis of the genus is presented. Shells and, as far as known, genitalia of 13 species are described and figured. Two species are described as new to science, i.e., A. vespa sp. nov. and A. enguriensis sp. nov. The diagnostic characteristics are summarized in a table. All locality records are listed and shown in distribution maps. Based on Bayesian, Maximum likelihood and Maximum parsimony analyses of mitochondrial and nuclear DNA sequences we revise the classification of the genus and suggest a division into three subgenera, Iliamneme, Acrotomina and Acrotoma s. str. The radiation of the Acrotoma species in the Pliocene and Quaternary might have been triggered by the orogenesis of the Greater Caucasus, which created habitats, such as canyons and alpine environments, to which the Acrotoma species adapted.www.zoobank.org/urn:lsid:zoobank.org:pub:6B14BE68-0E07-42EF-B08D-A75B2138B72F