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This study presents a complete species list of the door snails inhabiting Myanmar, updated to now include 33 taxa, and provides taxonomic notes together with a re-description of the shell, radula, and genitalia for 13 species and subspecies, including Oospira philippiana, the type species of the genus Oospira. The snails previously treated as subspecies or synonyms of Oospira gracilior and Oospira magna are reclassified and recognized as distinct species. The lectotype of Oospira insignis has been clarified and an illustration of the original type specimen provided. A long-overlooked species, Oospira andersoniana, has been collected and redescribed herein. Two new species from the limestone karsts in the Salween River Basin are introduced: Oospira luneainopsis Man & Panha, sp. nov. and Oospira zediopsis Man & Panha, sp. nov. A synoptic view of all clausiliid taxa known from Myanmar is presented along with taxonomic information and distributional records. Photographs of the type materials for all taxa are provided for further comparison or, if unavailable, photographs of the examined specimens or the original figure from the literature.

Many Nerita species live in warm-water environments, and they are some of the few organisms from the intertidal zone that can live in both freshwater and seawater. Previous comparative studies of the mitogenomes of Nerita species suggest that the genome rearrangements are very conservative. Generally, the species possess a set of similar mitochondrial gene arrangements, but nucleotide sequences can be used to elucidate phylogenetic relationships at various levels of divergence. Here, the mitogenomes of Nerita undata and Nerita balteata were sequenced and found to be 15,583 bp and 15,571 bp, respectively. The mitogenomes of both species contain 13 protein-coding genes, 22 tRNA genes, and two rRNA genes. The nucleotides of the two mitogenomes are highly similar, with the same gene composition and genomic organization as those present in other Nerita species. The tRNA secondary structures were different from those of other gastropods: trnS2 is not folded into typical secondary structures, and the dihydrouridine (DHU) arm simply forms a loop. The phylogenetic analysis showed that Neritimorpha is a sister group of Vetigastropoda and Caenogastropoda. Nerita balteata is a sister group of Nerita versicolor and Nerita undata, and all three species belong to Neritimorpha. This study contributes towards the comparative mitogenomic analysis of Neritidae and phylogenetic considerations among Neritimorpha species. The estimation of divergence time revealed that the two Nerita species were differentiated in the late Paleogene of the Cenozoic Era, and their evolution may be related to environmental changes.

Freshwater snails usually possess thin unadorned shells lacking structural components such as spines. Exceptions can be found on the high, well-watered islands of the South Pacific. Streams on these islands support a rich freshwater molluscan fauna with several nerite taxa (Neritimorpha: Neritidae) exhibiting extremely long dorsal spines. We sought to assess the defensive capacity of these structures for several co-occurring nerite genera on the Island of Ovalau, Fiji. Our overarching hypothesis was that spines confer a defensive advantage. We tested four predictions for eight common taxa: (i) predator “rich” habitats (the creek entrance) would be dominated by spine-bearing nerites, (ii) spine-bearing species should be smaller in size, (iii) nerites with spines would exhibit lower levels of shell damage and (iv) nerites with spines should invest less in their shells (i.e., their shells should be thinner). Most of these predictions received support. Spine-bearing species dominated the entrance to the creek and were smaller in size. Levels of shell damage were low overall, with 2 of the 3 spinose taxa exhibiting no shell damage, as did many of the nonspinose taxa. Finally, shells of spinose species were 25% thicker, demonstrating increased rather than decreased investment. Taken together, these findings suggest that the elaborate spines of Clithon spp. play a defensive role.

For the first time gastropods from the Besano Formation (Anisian/Ladinian boundary) are documented. The material was collected from three different outcrops at Monte San Giorgio (Southern Alps, Ticino, Switzerland). The taxa here described are Worthenia (Humiliworthenia)? aff. microstriata, Frederikella cf. cancellata, ?Trachynerita sp., ?Omphaloptycha sp. 1 and ?Omphaloptycha sp. 2. They represent the best preserved specimens of a larger collection and document the presence in this formation of the clades Vetigastropoda, Neritimorpha and Caenogastropoda that were widespread on the Alpine Triassic carbonate platforms. True benthic molluscs are very rarely documented in the Besano Formation, which is interpreted as intra-platform basin sediments deposited in usually anoxic condition. Small and juvenile gastropods could have been lived as pseudoplankton attached to floating algae or as free-swimming veliger planktotrophic larval stages. Accumulations of larval specimens suggest unfavorable living conditions with prevailing disturbance in the planktic realm or mass mortality events. However, larger gastropods more probably were washed in with sediments disturbed by slumping and turbidite currents along the basin edge or storm activity across the platform of the time equivalent Middle San Salvatore Dolomite.

Metallothioneins’ (MTs) biological function has been a matter of debate since their discovery. The importance to categorize these cysteine-rich proteins with high coordinating capacity into a specific group led to numerous classification proposals. We proposed a classification based on their metal-binding abilities, gradually sorting them from those with high selectivity towards Zn/Cd to those that are Cu-specific. However, the study of the NpeMT1 and NpeMT2isoforms of Nerita peloronta, has put a new perspective on this classification. N. peloronta has been chosen as a representative mollusk to elucidate the metal-binding abilities of Neritimorpha MTs, an order without any MTs characterized recently. Both isoforms have been recombinantly synthesized in cultures supplemented with ZnII, CdII, or CuII, and the purified metal–MT complexes have been thoroughly characterized by spectroscopic and spectrometric methods, leading to results that confirmed that Neritimorpha share Cd-selective MTs with Caenogastropoda and Heterobranchia, solving a so far unresolved question. NpeMTs show high coordinating preferences towards divalent metal ions, although one of them (NpeMT1) shares features with the so-called genuine Zn-thioneins, while the other (NpeMT2) exhibits a higher preference for Cd. The dissimilarities between the two isoforms let a window open to a new proposal of chemical MT classification.

Metallothioneins’ (MTs) biological function has been a matter of debate since their discovery. The importance to categorize these cysteine-rich proteins with high coordinating capacity into a specific group led to numerous classification proposals. We proposed a classification based on their metal-binding abilities, gradually sorting them from those with high selectivity towards Zn/Cd to those that are Cu-specific. However, the study of the NpeMT1 and NpeMT2isoforms of Nerita peloronta, has put a new perspective on this classification. N. peloronta has been chosen as a representative mollusk to elucidate the metal-binding abilities of Neritimorpha MTs, an order without any MTs characterized recently. Both isoforms have been recombinantly synthesized in cultures supplemented with ZnII, CdII, or CuII, and the purified metal–MT complexes have been thoroughly characterized by spectroscopic and spectrometric methods, leading to results that confirmed that Neritimorpha share Cd-selective MTs with Caenogastropoda and Heterobranchia, solving a so far unresolved question. NpeMTs show high coordinating preferences towards divalent metal ions, although one of them (NpeMT1) shares features with the so-called genuine Zn-thioneins, while the other (NpeMT2) exhibits a higher preference for Cd. The dissimilarities between the two isoforms let a window open to a new proposal of chemical MT classification.

The mitochondrial genome of the neritid snail Nerita (Heminerita) japonica (Mollusca: Neritimorpha) from Kumamoto, Japan was determined by whole-genome sequencing. This mitogenome is comprised of 13 protein-coding genes, 2 ribosomal RNA (12S and 16S) genes, and 22 transfer RNA genes, with the same gene order as in the other species of the family Neritidae. A likelihood-based phylogenetic reconstruction recovered the subgenus Heminerita (including N. japonica as its type and N. yoldii from China) as monophyletic and sister to a clade with four species of the subgenera Nerita and Theliostyla.

Neritidae is one of the most diverse families of Neritimorpha and possesses euryhaline properties. Members of this family usually live on tropical and subtropical coasts and are mainly gregarious. The phylogenetic relationships between several subclasses of Gastropoda have been controversial for many years. With an increase in the number of described species of Neritidae, the knowledge of the evolutionary relationships in this family has improved. In the present study, we sequenced four complete mitochondrial genomes from two genera (Clithon and Nerita) and compared them with available complete mitochondrial genomes of Neritidae. Gene order exhibited a highly conserved pattern among three genera in the Neritidae family. Our results improved the phylogenetic resolution within Neritidae, and more comprehensive taxonomic sampling of subclass Neritimorpha was proposed. Furthermore, we reconstructed the divergence among the main lineages of 19 Neritimorpha taxa under an uncorrelated relaxed molecular clock.

The molluscan fauna of southwestern Brazilian Amazonia is poorly known due to the lack of focused collection areas in the region since the early 20th century. The present study provides an inventory of the terrestrial gastropods from a forest fragment in the eastern Acre state, Brazil: the Humaitá Forest Reserve ("Reserva Florestal Humaitá"). Live specimens and empty shells were collected between August 2018 and January 2019. A total of 20 species were identified, distributed in 11 families. Most (13) of these species were recorded in Acre for the first time. Furthermore, we confirm the occurrence of Systrophia helicycloides (d'Orbigny, 1835) in Brazil, reiterate the synonymy of Plekocheilus pentadinus (d'Orbigny, 1835) with P. floccosus (Spix in Wagner, 1827), and synonymize Solaropsis peruviana Haas, 1951 with S. juruana Ihering, 1905.

Samples of terrestrial gastropods were collected year-round in seven caves in Presidente Olegário municipality, Minas Gerais state, southeastern Brazil, during several expeditions from 2012 to 2014. Twenty-four taxa (plus a single freshwater species), mainly stylommatophorans, were found in the material. The following species are reported for the first time for Minas Gerais state: Alcadia iheringi Wagner, 1910 and Helicina sordida King, 1831 (Helicinidae); Cecilioides consobrina (d’Orbigny, 1841) (Ferussaciidae); Entodina gionensis Morretes, 1940 and Scolodonta interrupta (Suter, 1900) (Scolodontidae); Megalobulimus sanctipauli (Ihering & Pilsbry, 1900) (Strophocheilidae); Drymaeus coarctatus (Pfeiffer, 1845) (Bulimulidae); Habroconus semenlini (Moricand, 1846) (Euconulidae); and Solaropsis aff. rosaria (Pfeiffer, 1849) (Solaropsidae). Furthermore, the species Drymaeus iracema (Simone, 2015) and Drymaeus terreus (Simone, 2015) are synonymized with Drymaeus coarctatus (L. Pfeiffer, 1845). Resumo: Amostras de gastrópodes terrestres foram coletadas em sete cavernas na região do município de Presidente Olegário, Minas Gerais, Brasil, durane múltiplas expedições ao longo dos anos de 2012 a 2014. Vinte-e-quatro táxons (mais uma única espécie dulciaquícola) foram encontrados, em sua maioria Stylommatophora. As seguintes espécies são aqui reportadas pela primeira vez para o estado de Minas Gerais: Alcadia iheringi Wagner, 1910 e Helicina sordida King, 1831 (Helicinidae); Cecilioides consobrina (d’Orbigny, 1841) (Ferussaciidae); Entodina gionensis Morretes, 1940 e Scolodonta interrupta (Suter, 1900) (Scolodontidae); Megalobulimus sanctipauli (Ihering & Pilsbry, 1900) (Strophocheilidae); Drymaeus coarctatus (Pfeiffer, 1845) (Bulimulidae); Habroconus semenlini (Moricand, 1846) (Euconulidae); e Solaropsis aff. rosaria (Pfeiffer, 1849) (Solaropsidae). Além disso, as espécies Drymaeus iracema (Simone, 2015) e Drymaeus terreus (Simone, 2015) são aqui sinonimizadas com Drymaeus coarctatus (L. Pfeiffer, 1845).

The Southwest Pacific represents an independent biogeographic province for deep-sea hydrothermal vent fauna. Different degrees of genetic connectivity among vent fields in Manus, North Fiji and Lau Basins have been reported for various molluscan and crustacean species, presumably reflecting their different levels of dispersal ability as swimming larvae. The present study investigates the population connectivity of the hydrothermal vent limpet Shinkailepas tollmanni (family Phenacolepadidae) in the Southwest Pacific. Our analyses using mitochondrial COI-gene sequences and shell morphometric traits suggest a panmictic population structure throughout its geographic and bathymetric ranges, spanning 4,000 km from the westernmost Manus Basin (151ºE; 1,300 m deep) to the easternmost Lau Basin (176ºE; 2,720 m). The measurements of its embryonic and larval shells demonstrate that the species hatches as a planktotrophic veliger larva with an embryonic shell diameter of 170-180 μm and settles at the vent environment with the larval shell diameter of 750-770 μm. This substantial growth as a feeding larva, ca. 80 times in volume, is comparable or even greater than those of confamilial species in the hydrothermal-vent and methane-seep environments in the Northwest Pacific and Atlantic Oceans. Large pigmented eyes in newly settled juveniles are another common feature in this and other phenacolepadids inhabiting the chemosynthetic environments. These results put together suggest that the larvae of S. tollmanni migrate vertically from deep-sea vents to surface waters to take advantages of richer food supplies and faster currents and stay pelagic for an extended period of time (> 1 year), as previously indicated for the confamilial species.

Prior to this study, few collections and records were made of the land snails in Cambodia and the historical taxa had never been reviewed. Herein a report on the land snail diversity based on specimens collected recently from karstic and non-karstic areas in southern Cambodia is provided. This checklist presents 36 species of land snails (two Neritimorpha, six Caenogastropoda, and 28 Heterobranchia). Illustrations and brief taxonomic notes/remarks are provided for every species. We also describedGeorrisa carinataSutcharit & Jirapatrasilp,sp. nov.based on some distinct shell morphological characters. Since the first descriptions during the colonial period in the nineteenth century, some land snail species (e.g.,Trichochloritis norodomiana,Durgella russeola,Anceyoconcha siamensis obesulacomb. nov.,Anceyoconcha chaudoensiscomb. nov., andSuccinea tenuis) have not been reported subsequently. This probably reflects a lack of knowledge concerning land snail biodiversity in this country. To our knowledge, this is the first comprehensive survey of land snails in southern Cambodia. A need for more field research and systematic revision of the land snails in this interesting region is also highlighted and demonstrated.

A new early middle Anisian (Bithynian) gastropod fauna is described from the Mahmudia quarry in North Dobrogea, Romania. At least 22 species are reported, nearly all of them from Tubiphytes-limestones which represent a microbial buildup. Twenty species from Tubiphytes-buildup limestones are characterized, ten of which in open nomenclature due to insufficient preservation. One genus (Murchisonietta gen. nov) and ten species are described as new. The new species are Euryalox applanatus, Rasatomaria multistriata, Sisenna inaequistriata, Worthenia (Worthenia) dobrogeana, Worthenia (Humiliworthenia) microstriata, Worthenia (Humiliworthenia) anisica, Marmolatella iordanae, Ladinaticella simionescui, Murchisonietta acuta and Coelostylina micropunctata. The present fauna is one of the most diverse Anisian gastropod faunas known to date and sheds light on the pattern of ongoing recovery from the end-Permian mass extinction event. It is dominated by Neritimorpha, Caenogastropoda and Pleurotomariida both in abundance and number of species. Diversity of Pleurotomariida strongly exceeds that of any known Early Triassic gastropod fauna and suggest a pronounced rebound of this group. High-spired smooth caenogastropods (Omphaloptycha and Polygyrina) are relatively abundant but for the most part not enough well preserved for species identification. The present gastropod fauna is part of a diverse invertebrate fauna probably comprising more than 50 species with gastropods being the most diverse followed by bivalves and brachiopods. They were microbial-buildup dwellers while metazoan frame-builders, especially corals, were still absent as consequence of the end-Permian mass extinction event. The Mahmudia fauna suggests that metazoan reefs recovered more slowly than benthic faunas.

Here is given an account, based on museum collections, of the fossil land and freshwater gastropods from Dischingen (Baden-Wurttemberg state, SW Germany). The fossiliferous outcrops belong to the Silvana Beds unit of the Upper Freshwater Molasse group, of late Early to early Middle Miocene age (Neogene European Land Mammal Zone MN 5). In total, 35 gastropod species were found (seven freshwater and 28 terrestrial), mainly belonging to Pulmonata, but also with representatives of Neritimorpha and Caenogastropoda. Seven species are reported here for the first time from Dischingen: Palaina diezi, Radix socialis, Granaria sp., Serrulina? sp., Discus pleuradrus, Punctum pumilio, and Oxyloma minima.

The red-blooded limpet ‘Shinkailepas’briandi(Neritimorpha: Phenacolepadidae) is one of the commonest gastropod species at deep-sea hydrothermal vents on the Mid-Atlantic Ridge (MAR). We investigated its population connectivity along MAR as the first such study for gastropods and explored the importance of larval migration for the distribution of vent-endemic animals. Our analyses, based on 1.3-kbp DNA sequences from the mitochondrial COI gene, showed a panmictic population throughout its geographic and bathymetric ranges that span from the northernmost and shallowest Menez Gwen vent field (38°N; 814–831 m depth) to the southernmost and deepest Ashadze field (13°N; 4090 m). Early development of this species is presumed to have a long pelagic duration as a planktotrophic larva; the hatchling with a shell diameter of 170–180 μm attains a constant settlement size of 706 ± 8 μm (mean ± SD). Retention of eye pigmentation in newly settled juveniles, along with the genetic panmixia, suggests that the hatched larva of ‘S.’briandimigrates vertically to the surface water, presumably to take advantage of richer food supplies and stronger currents for dispersal, as has been shown for confamilial species at hydrothermal vents and cold methane seeps.

The radula is the organ for mechanical food processing and an important autapomorphy of Mollusca. Its chitinous membrane, embedding small radular teeth, is moved by the set of muscles resulting in an interaction with the ingesta, tearing it and collecting loosened particles. Radulae and their teeth can be quite distinct in their morphology and had been of high research interest, but only a few studies have examined the basic functional principles of this organ, the movement and motion during feeding action. Here, the radular motion of 20 representative species, belonging to four major gastropod lineages (Vetigastropoda, Neritimorpha, Caenogastropoda and Heterobranchia) and Polyplacophora, were recorded and classified. Comparisons of the video footage with the scanning electron microscope (SEM) images of the radula resulted in the recognition of functional tooth rows and the correct position of the teeth during feeding. We identified six different types of radular movements, including rotations and bending of the radula itself. In each movement type, different structures act as counter bearings enabling the animals to grab and tear food., Summary: The radula is the food-processing organ in Mollusca and one key innovation. By employing high resolution video footage we documented distinct motion patterns while feeding and discovered that the animals can use the radula as counter bearings enabling the tearing of large food items.

The Bornean representatives of the genus Georissa (Hydrocenidae) have small, dextral, conical, calcareous shells consisting of ca. three teleoconch whorls. Our recent study on the Georissa of Malaysian Borneo has revealed high intra- and inter-specific variation in the “scaly” group (a group of species with striking scale-like surface sculpture). The present study on the “non-scaly” Georissa is the continuation of the species revision for the genus. The “non-scaly” species are also diverse in shell sculptures. This informal group comprises Georissa with subtle spiral and/or radial sculpture. The combination of detailed conchological assessment and molecular analyses provides clear distinctions for each of the species. Conchological, molecular, and biogeographic details are presented for 16 species of “non-scaly” Georissa. Three of these are new to science, namely Georissacorrugatasp. n., Georissainsulaesp. n., and Georissatrusmadisp. n.

Gunung Kidul is one of the districts in Yogyakarta that has a long coastline with coral substrate and white sand. One of the beaches with coral and rocky substrates is Indrayanti Beach. The rocky substrate provides unique characteristics for the life of marine organisms in the intertidal zone. Gastropods is one of the seven classes in the Mollusc phylum. Gastropods are found mostly in the sea and fresh water, especially in the intertidal zone of beach. The aim of this study is to identify gastropods that found in the Indrayanti intertidal zone and classifying each species. The method of this study is observation and sampling techniques with accidental sampling at low tide. Data analysis is descriptive based on morphological characteristics of each species. Based on the results of this study, there are 5 sub-classes, 9 orders, 2 Superfamily, 21 families, 35 genera and 72 species. Sub-classes found are heterobranchia, caenogastropods, neritimorpha, vestigastropods, and patelogastropods. The highest number of species found in the caenogastropod subclass was 32 species (44% of the total species) which were dominated by the order of neogastropod.

Gastropod mollusks are arguably the most diverse and abundant animals in the oceans, and are successful colonizers of terrestrial and freshwater environments. Here we resolve deep relationships between the five major gastropod lineages - Caenogastropoda, Heterobranchia, Neritimorpha, Patellogastropoda and Vetigastropoda - with highly congruent and supported phylogenomic analyses. We expand taxon sampling for underrepresented lineages with new transcriptomes, and conduct analyses accounting for the most pervasive sources of systematic errors in large datasets, namely compositional heterogeneity, site heterogeneity, heterotachy, variation in evolutionary rates among genes, matrix completeness and gene tree conflict. We find that vetigastropods and patellogastropods are sister taxa, and that neritimorphs are the sister group to caenogastropods and heterobranchs. With this topology, we reject the traditional Archaeogastropoda, which united neritimorphs, vetigastropods and patellogastropods, and is still used in the organization of collections of many natural history museums. Several traits related to development and life history support our molecular results. Importantly, the time of differentiation of the embryonic 4d cell (mesentoblast, responsible for mesoderm formation), differs between the two major clades, highlighting the degree of conservation and significance of development in the evolution of gastropods, as it is also known for spiralian animals more broadly.

Mannitol oxidase and polyol dehydrogenases are enzymes that convert polyalcohols into sugars. Mannitol oxidase was previously investigated in terrestrial snails and slugs, being also present in a few aquatic gastropods. However, the overall distribution of this enzyme in the Gastropoda was not known. Polyol dehydrogenases are also poorly studied in gastropods and other mollusks. In this study, polyalcohol oxidase and dehydrogenase activities were assayed in the digestive gland of 26 species of gastropods, representing the clades Patellogastropoda, Neritimorpha, Vetigastropoda, Caenogastropoda and Heterobranchia. Marine, freshwater and terrestrial species, including herbivores and carnivores were analyzed. Ultrastructural observations were undertake in species possessing mannitol oxidase, in order to investigate the correlation between this enzyme and the presence of tubular structures known to be associated with it. Mannitol oxidase activity was detected in the digestive gland of herbivores from the clades Caenogastropoda and Heterobranchia, but not in any carnivores or in herbivores from the clades Patellogastropoda, Neritimorpha and Vetigastropoda. In most of the species used in this study, dehydrogenase activities were detected using both D-mannitol and D-sorbitol as substrates. Nevertheless, in some carnivores these activities were not detected with both polyalcohols. Ultrastructural observations revealed tubular structures in digestive gland cells of some species having mannitol oxidase activity, but they were not observed in others. Based on our results, we suggest that mannitol oxidase first occurred in a herbivorous or omnivorous ancestor of Apogastropoda, the clade formed by caenogastropods and heterobranchs, being subsequently lost in those species that shifted towards a carnivorous diet. This study was supported by funds provided by the Abel Salazar Institute of Biomedical Sciences (ICBAS) of the University of Porto (Portugal), and by Fundação para a Ciência e Tecnologia (FCT) through the strategic project UID/ MAR/04292/2013 granted to MARE. The funder had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.

Tiny, circular objects preserved in the form of rims have been detected on an Early Cretaceous (early Aptian) ammonite from Daghestan, Russia. They are preserved on the body chamber portion of the mould, where they occur either as isolated rims or, more commonly, as structures closely neighbouring with each other. Comparisons with similar Recent and fossil structures indicate that they are remnants of gastropod egg capsules preserved as attachment bases, and most probably were produced by neritimorph gastropods. The egg capsules were deposited within an empty ammonite body chamber where the gastropods found a site sheltered against potential scavengers and predators. The occurrence of only attachment bases indicates, that the egg capsules may have hatched. Although they are preserved in the form of iron oxides, during fossilization the originally organic egg capsules underwent pyritization followed by later weathering. This is the first record of this kind from the Lower Cretaceous.

The identity of Helicina mediana Gassies, 1870 was clarified by the study of the type material and the selection of a lectotype. It is currently classified as Sturanya mediana (Gassies, 1870) and replaces the temporarily applied name Sturanya novaecaledoniae (Baird, 1873) for the medium sized helicinid species widely spread in the north-eastern part of mainland New Caledonia and the adjacent Îles Belep. The latter name and Helicina nehoueensis Hartman 1889 are synonyms of S. mediana.

Successful responses to the multifarious challenges of controlling water balance are critical for snails' survival in the great diversity of habitats they occupy. Advances are being made in understanding how such challenges are approached at the molecular level, including through the study of aquaporins, which are proteins functioning to facilitate the passage of water and other small molecules across cellular membranes. Deduced aquaporin amino acid sequences from partial genomic assemblies of three neritimorph species were added to available gastropod data and sequences from other taxa to make a phylogenetic classification of these proteins using maximum likelihood and Bayesian analyses. We identified ten groups, designated as G1 to G10, containing sequences from multiple major gastropod lineages. At least six of the groups appear to be encoded by multiple genes within at least some species. Five weakly-associated sequences from Neritimorpha were not allocated to a group. The designated groups G1, G2, G3, G4, G5 and G7 (previously defined as Malacoglyceroporins) formed clades containing only gastropod sequences and were strongly supported by Bayesian inference. G1, G2, G3 and G5 were also strongly supported by maximum likelihood analyses. Group G6 (previously defined as Malacoaquaporins)was included with sequences from the oyster, Crassostrea gigas in a strongly supported clade. Groups G8 and G9 included only gastropod sequences but were not strongly supported. Groups G8 and G10 were designated to include all the gastropod sequences belonging respectively to strongly-supported clades including human aquaglyceroporins and aquaammoniaporins. Most groups have been found in a wide range of gastropod lineages but all identified representatives of group G7 belong to Apogastropoda whereas G2 is known only from Patellogastropoda and Neritimorpha.

The complete mitochondrial genome of Nerita albicilla was determined. The mitochondrial genome is 15,314 bp in length, comprising two ribosomal RNA (12S and 16S) genes, 13 protein-coding genes, and...

In the present study, we determined the mitochondrial DNA (mtDNA) sequence of three Neritas, Nerita versicolor, Nerita tessellata, and Nerita fulgurans. We present an analysis of the features of their gene content and genome organization and compare these within the genus Nerita, and among the main gastropod groups. The new sequences were used in a phylogenetic analysis including all available gastropod mitochondrial genomes. Genomic lengths were quite conserved, being 15,866bp for N. versicolor, 15,741bp for N. tessellata and 15,343bp for N. fulgurans. Intergenic regions were generally short; genes are transcribed from both strands and have a nucleotide composition high in A and T. The high similarity in nucleotide content of the different sequences, gene composition, as well as an identical genomic organization among the Nerita species compared in this study, indicates a high degree of conservation within this diverse genus. Values ​​of Ka/Ks of the 13 protein coding genes (PCGs) of Nerita species ranged from 0 to 0.18, and suggested different selection pressures in gene sequences. Bayesian phylogenetic analyses using concatenated DNA sequences of the 13 PCGs and the two rRNAs, and of amino acid sequences strongly supported Neritimorpha and Vetigastropoda as sister groups.

Body size during larval development is one of the most important attributes of aquatic animals. The optimal size for changing form or habitat may vary according to ecological traits of species, while phylogenetic constraints also play a significant role. The major goal of this study is to reveal the patterns in the settlement size of planktotrophic larvae in an archaic gastropod superorder Neritimorpha. We take advantage of the fact that size at various ontogenetic stages of neritimorphs can be rigorously estimated by measuring features of the adult opercula. This unique feature of neritimorphs has allowed us to generate the largest data set so far on larval settlement sizes within a group of marine invertebrates that recruit into very different post-metamorphic habitats. Eighty-eight species that represent most extant genera from rocky shores, seagrass beds, mangroves, estuaries, streams, submarine caves, deep-sea vents or seeps showed negligible intraspecific variation and considerable interspecific differences in settlement size, particularly between genera or families. Settlement size is determined primarily by phylogenetic constraints, while parallel evolution toward smaller sizes was shown to occur exclusively in four independent clades (two living and two extinct) of amphidromous snails with a marine larval period followed by a limnic adult phase. The smaller settlement size may possibly reduce the risk of being wafted away from the estuaries of their natal streams through less time achieving metamorphic competence, while ability to make occasional long-distance trips is retained by the presence of a sufficiently long delay period. This delay period also seems to obscure the possible correlation between settlement size and geographic distribution range of neritimorph species, both fully marine and amphidromous.

The egg capsules of neritid snails (Gastropoda: Neritimorpha) are made of a tough inner organic layer and an outer calcareous layer for further reinforcement. Such hard capsules are less susceptible to mechanical damage compared with soft or thin capsules of other gastropods, and there have been only a few reports of predation on neritid eggs. Here, we describe drilling predation on the reinforced egg capsule of Nerita japonica by Reishia clavigera (Caenogastropoda: Muricidae) on a high intertidal rocky shore in central Japan. The muricid makes a small, elliptical drill hole in the capsule lid by the use of both an accessory boring organ (ABO) and a radula, as it does in mollusc and barnacle shells. The etching around the hole by the acid secretion from the ABO, which can be observed under a dissecting microscope and by scanning electron microscopy, should provide an effective way to identify muricid predation on neritid eggs, probably not only by R. clavigera but also by other species in the warm temperate to tropical regions all over the world. Reishia clavigera made drill holes in c. 14% of the capsules, making it possibly the main cause of mortality for neritid eggs at the study site. However, this mortality rate is lower than those previously reported for caenogastropod eggs with softer capsules. All penetrated capsules of N. japonica were white (rather than brownish) and newly laid, suggesting that their acid-resistant organic layer had yet to harden at the time of radular rasping. The thick organic layer and calcareous coating are an effective armour, not only against the usual type of grazing but also against more sophisticated predation involving the chemical and mechanical boring of Muricidae.

The limpet form – a cap-shaped or slipper-shaped univalved shell – convergently evolved in many gastropod lineages, but questions remain about when, how often, and under which circumstances it originated. Except for some predation-resistant limpets in shallow-water marine environments, limpets are not well adapted to intense competition and predation, leading to the prediction that they originated in refugial habitats where exposure to predators and competitors is low. A survey of fossil and living limpets indicates that the limpet form evolved independently in at least 54 lineages, with particularly frequent origins in early-diverging gastropod clades, as well as in Neritimorpha and Heterobranchia. There are at least 14 origins in freshwater and 10 in the deep sea, with known times ranging from the Cambrian to the Neogene. Shallow-water limpets are most diverse at mid-latitudes; predation-resistant taxa are rare in cold water and absent in freshwater. These patterns contrast with the mainly Late Cretaceous and Caenozoic warm-water origins of features such as the labral tooth, enveloped shell, varices, and burrowing-enhancing sculpture that confer defensive and competitive benefits on molluscs.

Vetigastropod and neritimorph species from Lower Bajocian (Middle Jurassic) sediments of south-western Luxembourg are described. Eighteen species are recognized. Two new genera – Szabotomaria gen. nov. and Fabercapulus gen. nov. – and four new species – Szabotomaria ziqquratiformis sp. nov., Colpomphalus thuyi sp. nov., Colpomphalus tigratus sp. nov. and Fabercapulus semisculptus sp. nov. – are erected. These species, together with other species recently described from the same strata, are part of a diverse assemblage consisting of 32 species belonging to 14 genera in six families and five superfamilies, representing the richest vetigastropod-neritimorph fauna currently known from Bajocian and sub-coeval deposits of western Europe. An analysis of the palaeoecological and palaeobiogeographical relationships of this fauna and those from other areas of the western European shelf was performed using cluster analysis and by comparison of the respective taxonomic structures. The former detected a major cluster ...

Larval dispersal significantly contributes to the geographic distribution, population dynamics, and evolutionary processes of animals endemic to deep-sea hydrothermal vents. Little is known as to the extent that their larvae migrate vertically to shallower waters and experience stronger currents and richer food supplies. Here, we first provide evidence from early life-history traits and population genetics for the surface dispersal of a vent species. Planktotrophic larvae of a red blood limpet, Shinkailepas myojinensis (Gastropoda: Neritimorpha: Phenacolepadidae), were cultured to observe their swimming behavior and to evaluate the effects of temperature on survival and growth. In addition, the population structure was analyzed based on 1.2-kbp mitochondrial DNA sequences from 77 specimens that cover the geographic and bathymetric distributions of the species (northwest Pacific, 442-1,227 m in depth). Hatched larvae constantly swam upward at 16.6-44.2 mm/min depending on temperature. Vertical migration from hydrothermal vents to the surface, calculated to take ~4-43 d, is attainable given their lengthy survival time without feeding. Fed larvae best survived and grew at 25°C (followed by 20°C), which approximates the sea surface temperature in the geographic range of the species. Little or no growth was observed at the temperature of the vent habitat where adult limpets occur (≤15°C). Population genetic analyses showed no differentiation among localities that are

Despite the extraordinary morphological and ecological diversity of Neritimorpha, few studies have focused on the phylogenetic relationships of this lineage of gastropods, which includes four extant superfamilies: Neritopsoidea, Hydrocenoidea, Helicinoidea, and Neritoidea. Here, the nucleotide sequences of the complete mitochondrial genomes of Georissa bangueyensis (Hydrocenoidea), Neritina usnea (Neritoidea), and Pleuropoma jana (Helicinoidea) and the nearly complete mt genomes of Titiscania sp. (Neritopsoidea) and Theodoxus fluviatilis (Neritoidea) were determined. Phylogenetic reconstructions using probabilistic methods were based on mitochondrial (13 protein coding genes and two ribosomal rRNA genes), nuclear (partial 28S rRNA, 18S rRNA, actin, and histone H3 genes) and combined sequence data sets. All phylogenetic analyses except one converged on a single, highly supported tree in which Neritopsoidea was recovered as the sister group of a clade including Helicinoidea as the sister group of Hydrocenoidea and Neritoidea. This topology agrees with the fossil record and supports at least three independent invasions of land by neritimorph snails. The mitochondrial genomes of Titiscania sp., G. bangueyensis, N. usnea, and T. fluviatilis share the same gene organization previously described for Nerita mt genomes whereas that of P. jana has undergone major rearrangements. We sequenced about half of the mitochondrial genome of another species of Helicinoidea, Viana regina, and confirmed that this species shares the highly derived gene order of P. jana., This work was supported by the Spanish Ministry of Science and Innovation (CGL2010-18216 and CGL2013-45211-C2-2-P to RZ; BES-2011-051469 to JEU)

Two of the three major gastropod clades with feeding larvae are sister groups and larval morphogenesis for members of these clades, the Caenogastropoda and Heterobranchia, has been well studied. The third clade, the Neritimorpha, has an unstable phylogenetic position and little is known about development of their planktotrophic larvae. Information about larval morphology of neritimorphs and resolution of their controversial phylogenetic placement is critically important for understanding evolution of larval feeding within the Gastropoda. We describe larval morphogenesis to metamorphic competence for laboratory-reared larvae of Nerita melanotragus (Smith, 1884) (Neritimorpha: Neritidae). Preliminary observations suggest that prehatch larvae are capable of delayed hatching, possibly by entering a diapause state. Our description of larval morphogenesis, as based on tissue sections for light and transmission electron microscopy, scanning electron microscopy, three-dimensional-reconstructions of sectioned tissue, and labeling of muscles with fluorphore-tagged phalloidin, revealed four features that are unprecedented among both feeding and nonfeeding gastropod larvae. Larvae of N. melanotragus have muscles on the left and right side that both meet current criteria of a larval retractor muscle; shell-anchored muscles with oblique striations that project inside the visceral nerve loop to insert mainly on the velar lobes. They also have left and right digestive glands of similar size and a left and right hypobranchial gland. A larval "heart" is absent, but water circulation through the mantle cavity may be facilitated by large circular orifices, lined by patches of motile cilia, leading in and out of the mantle cavity. Comparison of larval traits among all three groups of gastropods with feeding larvae indicates that larvae of N. melanotragus have many unique characteristics, but they show more similarities to caenogastropod than to heterobranch larvae. These results are a significant step toward the goal of identifying primitive versus derived larval traits among feeding gastropod larvae.

SUMMARY Early and middle Paleozoic gastropod protoconchs generally differ strongly from their corresponding adult morphologies, that is, most known protoconchs are smooth and openly coiled, whereas the majority of adult shells are ornamented and tightly coiled. In contrast, larval and adult shells of late Paleozoic gastropods with planktotrophic larval development (Caenogastropoda, Neritimorpha) commonly resemble each other in shape and principle ornamentation. This is surprising because habitat and mode of life of planktonic larvae and benthic adults differ strongly from each other. Generally, late Paleozoic to Recent protoconchs are tightly coiled. This modern type of larval shell resembles the adult shell morphology and was obviously predisplaced onto the larval stage during the middle Paleozoic. The oldest known planktonic-armored (strongly ornamented) larval shells are known from the late Paleozoic. However, smooth larval shells are also common among the studied late Paleozoic gastropods. The appearance of larval armor at the beginning of the late Paleozoic could reflect an increase of predation pressure in the plankton. Although there are counter examples in which larval and adult shell morphology differ strongly from each other, there is statistical evidence for a heterochronic predisplacement of adult characters onto the larval stage. Larval and adult shells are built in the same way, by accretionary secretion at the mantle edge. It is likely that the same underlying gene expression is responsible for that. If so, similarities of larval and adult shell may be explained by gene sharing, whereas differences may be due to different (planktic vs. benthic life) epigenetic patterns.

This is the first report of the mitochondrial gene order and almost-complete DNA sequence of a representative of the Neritimorpha, the highest-ranking gastropod clade lacking such data. Mitochondrial gene order in Nerita is largely plesiomorphic. Its only difference from the cephalopod Octopus vulgaris is a tRNA transposition shared by Vetigastropoda and Caenogastropoda. Genome arrangements were not informative enough to resolve the evolutionary relationships of Neritimorpha, Vetigastropoda and Caenogastropoda. The sister-group taxon of Neritimorpha varied in sequence-based analyses. Some suggested that Neritimorpha is the sister group of Caenogastropoda plus Heterobranchia and some that Neritimorpha and Caenogastropoda are sister groups. No analysis significantly supported the hypothesis that Vetigastroda is more closely related to Caenogastropoda than is Neritimorpha.

Helicina variabilis Wagner, 1827 (Neritimorpha, Helicinidae) is redescribed based on a sample collected in Nanuque, northern Minas Gerais, Brazil. The species description, previously based only on the shell, is expanded to the phenotypic features. The study revealed absorption of the internal shell whorls; a diaphragm muscle connected to the floor of the pallial cavity; a monoaulic pallial oviduct, with the female genital aperture inside the anal aperture, and the lack of a seminal receptacle and provaginal sac; and the pleural ganglia of the nerve ring connected with each other. The significance of these findings is discussed in the light of current taxonomic and phylogenetic knowledge. Helicina variabilis Wagner, 1827 (Neritimorpha, Helicinidae) is redescribed based on a sample collected in Nanuque, northern Minas Gerais, Brazil. The species description, previously based only on the shell, is expanded to the phenotypic features. The study revealed absorption of the internal shell whorls; a diaphragm muscle connected to the floor of the pallial cavity; a monoaulic pallial oviduct, with the female genital aperture inside the anal aperture, and the lack of a seminal receptacle and provaginal sac; and the pleural ganglia of the nerve ring connected with each other. The significance of these findings is discussed in the light of current taxonomic and phylogenetic knowledge.

The phylogenetic relationships among the “archaeogastropod” clades Patellogastropoda, Vetigastropoda, Neritimorpha, and Neomphalina are uncertain; the phylogenetic placement of these clades varies across different analyses, and particularly among those using morphological characteristics and those relying on molecular data. This study explores the relationships among these groups using a combined analysis with seven molecular loci (18S rRNA, 28S rRNA, histone H3, 16S rRNA, cytochrome c oxidase subunit I [COI], myosin heavy-chain type II, and elongation factor-1α [EF-1α]) sequenced for 31 ingroup taxa and eight outgroup taxa. The deep evolutionary splits among these groups have made resolution of stable relationships difficult, and so EF-1α and myosin are used in an attempt to re-examine these ancient radiation events. Three phylogenetic analyses were performed utilizing all seven genes: a single-step direct optimization analysis using parsimony, and two-step approaches using parsimony and maximum likelihood. A single-step direct optimization parsimony analysis was also performed using only five molecular loci (18S rRNA, 28S rRNA, histone H3, 16S rRNA, and COI) in order to determine the utility of EF-1α and myosin in resolving deep relationships. In the likelihood and POY optimal phylogenetic analyses, Gastropoda, Caenogastropoda, Neritimorpha, Neomphalina, and Patellogastropoda were monophyletic. Additionally, Neomphalina and Pleurotomariidae fell outside the remaining vetigastropods, indicating the need for further investigation into the relationship of these groups with other gastropods.