The mitochondrial genome of Ifremeria nautilei and the phylogenetic position of the enigmatic deep-sea Abyssochrysoidea (Mollusca: Gastropoda)

The complete nucleotide sequence of the mitochondrial (mt) genome of the deep-sea vent snail Ifremeria nautilei (Gastropoda: Abyssochrysoidea) was determined. The double stranded circular molecule is 15,664 pb in length and encodes for the typical 37 metazoan mitochondrial genes. The gene arrangement of the Ifremeria mt genome is most similar to genome organization of caenogastropods and differs only on the relative position of the trnW gene. The deduced amino acid sequences of the mt protein coding genes of Ifremeria mt genome were aligned with orthologous sequences from representatives of the main lineages of gastropods and phylogenetic relationships were inferred. The reconstructed phylogeny supports that Ifremeria belongs to Caenogastropoda and that it is closely related to hypsogastropod superfamilies. Results were compared with a reconstructed nuclear-based phylogeny. Moreover, a relaxed molecular-clock timetree calibrated with fossils dated the divergence of Abyssochrysoidea in the Late Jurassic–Early Cretaceous indicating a relatively modern colonization of deep-sea environments by these snails.     

The anatomy of Addisonia (Mollusca,Gastropoda)

Summary The organization of Addisonia lateralis (Requien, 1848) and A. brophyi McLean, 1985 is described. Addisonia species have a thin, asymmetrical, cup-like shell and a very simple shell muscle. Eyes, oral lappets and epipodial tentacles are absent and the right cephalic tentacle is also used as a copulatory organ. Most characteristic is the enormously developed gill which is enlarged into the right subpallial cavity. It is composed of about 30 leaflets with skeletal rods and its epithelia are uniquely arranged. The heart is large and the single auricle is situated anteriorly left. There are two kidneys: the left is small, while the right forms large coelomic cavities and has no connection with the pericardium or the hermaphroditic genital system. Testis and ovary are separate: both have a simple duct proper (vas deferens, oviduct). They are connected to the copulatory organ by an open seminal groove; a small receptaculum is present. The mouth opening is typically triangular, with no jaws or subradular sense organs. Addisonia possesses tuft-like salivary glands, a radula diverticulum and distinct, tubular oesophageal glands. The oesophagus itself is simple. The radula and the posterior alimentary tract are unique; the stomach is completely reduced and the intestine forms a pseudostomach. The streptoneurous, hyoathroid nervous system has pedal cords with three commissures. The visceral loop is also cord-like. A single (left) osphradium is present and the small statocysts have several statocones.The peculiarities and unique combination of primitive and advanced characters in Addisonia reflect a highly enigmatic organization among the Archaeogastropoda. Possible relationships to other archaeogastropod groups are discussed.     

A comparative ultrastructural investigation of the cephalic sensory organs in Opisthobranchia (Mollusca, Gastropoda)

Cephalic sensory organs (CSOs) are specialised structures in the head region of adult Opisthobranchia involved in perception of different stimuli. The gross morphology of these organs differs considerably among taxa. The current study aims at describing the cellular morphology of the CSOs in order to reveal cellular patterns, especially of sensory epithelia, common for opisthobranchs. Transmission electron microscopy was used to characterise the fine structure of the organs and to compare the CSOs of four different opisthobranch species. The cellular composition of the sensory system is conserved among taxa. The epidermal cells in sensory regions are always columnar and ciliated cells are frequently apparent. The sensory cells are primary receptors arranged in subepidermal cell clusters. They extend dendrites which penetrate the epithelium and reach the surface. Some of the dendrites bear cilia, whereas others only build a small protuberance. Processing of sensory information takes place in the peripheral glomeruli of all species. Moreover, few taxa possess additional peripheral ganglia at the base of their CSOs. The results of the present study might support other investigations indicating that the posterior CSOs are primarily involved in distance chemoreception, whereas the anterior CSOs might be used for contact chemoreception and mechanoreception.     

Tissue Hemoglobins of Gastropoda (Mollusca)

The distribution and content of tissue hemoglobins in the radular muscle, subradular cartilages, myocardium, and nerve tissue of gastropods are analyzed.     

The Vetigastropoda and the systematics of streptoneurous Gastropoda (Mollusca)

The status of the Vetigastropoda, created 1980 to unite zeugobranch and trochoid archaeo-gastropods, is reconsidered and confirmed in the light of recent investigations. The most important synapomorphic characters of the Vetigastropoda are the ctenidial sense organs (so-called bursicles), the epipodial sense organs, and the special structure of the oesophagus. The conditions of the archaeogastropod oesophagus are comparatively described.
An examination of the relationships of the Vetigastropoda establishes the archaeogastropod nature of the Architaenioglossa. It also demonstrates the monophyletic origin of all higher gastropods, including the Caenogastropoda (s.str.) as well as the Pentaganglionata (= Euthy-neura). The Campanilidae, Valvatidae, Rissoellidae, Omalogyridae, and the Allogastropoda (= Architectonicoidea and Pyramidelloidea) share characteristics with both of the above-mentioned groups, but cannot be classified within any of them. These groups are regarded as subsequent offshoots between the caenogastropod and the pentaganglionate level of organization. The level of Archaeogastropoda is redefined by the hypoathroid/dystenoid nervous system and simple tentacle nerves; the remaining Streptoneura are classified as Apogastropoda with epiathroid nervous system, parapedal commissure, and bifurcated tentacle nerves.     

Polygyrid relations: a phylogenetic analysis of 17 subfamilies of land snails (Mollusca: Gastropoda: Stylommatophora)

The family of polygyrid land snails in North America is significant for its sympatric shell convergences, diversity of mating systems and complex zoogeography; its monophyly and its relation to other families has long been questionable. Cladistic analysis has been performed on one representative each of 17 subfamilies, including all three polygyrid subfamilies and one or more subfamilies each from all ten of the families that have been proposed as the polygyrid sister groups. Eighteen anatomical synapomorphies are used, of which eight are newly discovered, four are differently assessed from previous studies, and six are traditional. The resulting strict consensus tree of alternative maximum-parsimony cladograms is: (Acavidae (Ammonitellidae Corillidae ((Discidae Oreohelicidae) (Helminthoglyptidae Bradybaenidae Polygyridae (Thysanophoridae (Camaenidae Sagdidae)))))).
According to this working hypothesis, the Polygyridae are monophyletic, and their sister group remains unresolved, although the Acavidae, Ammonitellidae, Corillidae, Discidae and Oreohelicidae can be ruled out. Of the five classifications of stylommatophoran families that have been proposed in the past 12 years, the consensus tree is closest to that of Nordsieck. For future morphological work, three regions are recommended as potentially rich in unused phylogenetic information: the fertilization pouch-seminal receptacle complex, the ureter at the pneumostome and the ventral-chain ganglia. Simultaneous dissection, wkh side-by-side comparisons, is recommended over other methods for detecting homologies in land snails. Molecular characters should be exploited, because of the scarcity and the frequent homoplasy of morphological characters.     

Basal chromodorid sperm ultrastructure (Nudibranchia, Gastropoda, Mollusca)

The relationship between three genera considered basal in the Chromodorididae (Cadlina, Tyrinna, Cadlinella) has not yet been resolved by traditional morphological means. Here we examined the sperm ultrastructure of Tyrinna nobilis, Tyrinna evelinae, Cadlina flavomaculata and Cadlina cf. nigrobranchiata, with the expectation of finding phylogenetically informative characters. No Tyrinna or Cadlina species showed sperm similarities to Cadlinella. Both Cadlina species and Tyrinna nobilis (but not T. evelinae) exhibited coarse striations in the acrosomal pedestal. The putative fibers that occurred between the coarse striations of the pedestal are condensed into a layer in Cadlina and Tyrinna, but not in other species that also have coarse striations (Gymnodoris), and may constitute evidence for a close relationship. Tyrinna evelinae possessed fine acrosomal striations, which was shared with other Chromodorididae, Actinocyclidae and the cryptobranchs Rostanga and Aphelodoris. We also examined the sperm ultrastructure of ‘Chromodoris’ ambiguus, an animal which has shown molecular affinities to species of Cadlina, and not Chromodoris. The sperm of ‘C.’ ambiguus did not exhibit the typical Cadlina characteristics, but also showed important differences to other investigated Chromodoris species.     

Studies on Priotrochus obscurus and the systematic position of Priotrochus (Mollusca: Gastropoda: Trochidae)

The systematic status of Priotrochus Fischer, 1879 and the taxonomy of its type species, Trochus obscurus Wood, 1828, are discussed. Although previously regarded as a subgenus of Monilea , observations on the radula and external anatomy reveal little similarity with Monilea and the Umboniinae as a whole. Greater similarity exists with members of the Monodontinae and a position, at generic level, in that subfamily is suggested. The extinct Trochus ponsonbyi Sowerby, 1888, is maintained as a subspecies of Priotrochus obscurus .     

Dart formation in Helix aspersa (Mollusca,Gastropoda)

Summary Dart formation in Helix aspersa has been investigated by SEM of isolated darts at progressive stages in their development, and by histology of dart sacs at the same times. Dart formation begins at the tip of a tubercle where a small group of epithelial cells secrete an organic material filling a small CaCO₃ cone that is the first mineralized part of the shaft. Subsequent secretory activity by an increasing area of the tubercle epithelium results in an increase in the diameter and anterior lengthening of the shaft. Continued secretion by the tubercle and dart sac epithelium produces the flare and finally the corona. A pattern of deposition is also evident in the fine structure of the mineral. In the shaft and vanes there is an inner layer of spherulitic prismatic structure which is covered by a layer of irregular patches of simple prismatic structure. The outermost layer of the shaft and vanes has a continuous simple prismatic structure. Two layers are present in the flare, an inner granular amorphous layer and an outer spherulitic prismatic layer. The corona consists of a single rarefied prismatic layer. A mechanism of dart formation is suggested that involves two types of organic matrix, calcifying and non-calcifying. Measurements of the calcium content of darts, dart sacs, and collars indicate that the hemolymph is the probable source of calcium for the dart.     

Systematic position of the pelagic Thecosomata and Gymnosomata within Opisthobranchia (Mollusca, Gastropoda) – revival of the Pteropoda

The complete 18S (SSU) rRNA as well partial 28S (LSU) rRNA and partial mitochondrial COI sequences have been used to reconstruct the phylogenetic relationships within Opisthobranchia with special focus on the pelagic orders Thecosomata and Gymnosomata. Maximum parsimony, maximum likelihood, distance as well as Bayesian analysis of a combined dataset of the three genes reveals that Thecosomata and Gymnosomata are sister groups and together are closely related to Anaspidea. Possible sister taxon to Thecosomata, Gymnosomata and Anaspidea is Cephalaspidea s. str . Analysis of a taxon-extended dataset of partial 28S sequences supported a basal position of Limacina within Euthecosomata. Within Cavolinidae, Creseis is basal to the other taxa. Other phylogenetic implications from the present results are also discussed. Investigation of the morphology and histology of Thecosomata and Gymnosomata as well as several other opisthobranch taxa helped to identify autapomorphies for Thecosomata and Gymnosomata as well as apomorphies for the clades including these taxa.     

Variations in the retinal designs of pulmonate snails (Mollusca, Gastropoda): squaring phylogenetic background and ecophysiological needs (I)

Abstract. The eyes of aquatic pulmonates differ from those of terrestrial pulmonates; the latter, in species such as Cepaea nemoralis and Trichia hispida , possess conventional, cup-shaped retinas, but the aquatic species Lymnaea stagnalis, Radix peregra, Physa fontinalis , and Planorbarius corneus have retinas that are partitioned into dorsal and ventral depressions ("pits"). The pits are separated by an internal ridge, called the "crest", and on account of their pigmentation can be seen in vivo . The dominant cellular components of the retinae of terrestrial as well as aquatic snails are pigmented cells and microvillar photoreceptors, the latter occurring in two morphologically distinct types (I and II). Aquatic snails with preferences for shallow water possess eyes with both type I and type II photoreceptive cells, but Pl. corneus , an inhabitant of deeper water, only has type-I receptors, supporting an earlier finding that type I cells represent dim- and type II cells bright-light receptors. On the basis of histological and optical comparisons, we conclude that the eyes of L. stagnalis and R. peregra , species that are known to escape and seek temporary refuge above the water surface, are well adapted to function in water as well as air, but that the eyes of P. fontinalis and Pl. corneus are less modified from those of their terrestrial ancestors.     

Fixed focal-length optics in pulmonate snails (Mollusca, Gastropoda): squaring phylogenetic background and ecophysiological needs (II)

Abstract. Our results suggest that freshwater pulmonates like Lymnaea stagnalis, Radix peregra, Physa fontinalis , and Planorbarius corneus have inherited from their terrestrial ancestors eyes with a spherical, immobile lens with fixed focal-length optics. Unable to change the dioptric apparatus to form an image under water, modifications to the retina had to occur if sharp vision was required. Computer-assisted calculations and 3-D eye reconstructions demonstrate that the photoreceptors in the deeper, ventral pit are in a position to perceive focused images under water. Vision in air, however, would favour photoreceptive cells located in the shallower, dorsal pit. On the basis of histological, ethological, and optical comparisons, we conclude that the eyes in L. stagnalis and R. peregra , species that are known to escape and seek temporary refuge above the water surface, are well adapted to function in water as well as air, but that the eyes in P. fontinalis and Pl. corneus are less modified from those of their terrestrial ancestors. We also conclude that good resolving power may be of greater importance in the aquatic pulmonates than the terrestrial species, since the former have to locate thin, vertical stems of reeds and sedges to ascend in order to reach the surface to breathe.     

An Ultrastructural Study of Basommatophoran Spermatozoa (Mollusca, Gastropoda)

Spermatozoa of the basommatophoran pulmonate families Ellobiidae ( Ophicardelus ornatus Ferussac), Amphibolidae ( Salinator fragilis Lamarck, Salinator solida von Martens), Siphonariidae ( Siphonaria funiculata Reeve) and Lymnaeidae ( Lymnaea lessoni (Deshayes)) were studied using transmission electron microscopy. Spermatozoa of all species, like most euthyneuran spermatozoa, possess ( 1 ) an acrosome composed of an apical vesicle and acrosomal pedestal (many differences between families), (2) a helically keeled, posteriorly invaginated nucleus, ( 3 ) a midpiece composed of paracrystalline and matrix materials and a variable number of incorporated glycogen-filled helices (one in Salinator and Siphonaria , two or three in Lymnaea , three in Ophicardelus ) and (4) an axoneme associated with coarse fibres (periodically banded in neck region) and rows of intra-axonemal granules. The wide diversity of spermatozoon structure in the species studied, in particular the midpiece structure of Siphonaria (which resembles closely that of certain opisthobranchs) indicates that the Basommatophora may not represent a valid taxonomic unit. A comparison of basommatophoran sperm with other euthyneurans and with euspermatozoa of prosobranchs is given.     

The Vermetidae of the Gulf of Kachchh,western coast of India (Mollusca,Gastropoda)

Coral reefs are often termed underwater wonderlands due to the presence of an incredible biodiversity including numerous invertebrates and vertebrates. Among the dense population of benthic and bottom-dwelling inhabitants of the reef, many significant species remain hidden or neglected by researchers. One such example is the vermetids, a unique group of marine gastropods. The present study attempts for the first time to assess the density and identify preferred reef substrates in the Gulf of Kachchh, state of Gujarat, on the western coast of India. A total of three species of the family Vermetidae were recorded during the study and their substrate preferences identified.     

Towards a monophyletic genus Albinaria (Gastropoda, Pulmonata): the first molecular study into the phylogenetic position of eastern Albinaria species

Recent molecular phylogenetic studies of the terrestrial snail genus Albinaria have caused a radical reassessment of its taxonomy. These studies, however, were limited to western species. This paper examines mitochondrial 12S sequences and nuclear ITS1 & 2 sequences of both eastern and western species, and demonstrates that Albinaria, in its most recent definition, is not monophyletic. Both molecular datasets place ' Albinaria ' hedenborgi from Lebanon in a well-supported clade with species of the genus Cristataria , distributed south-east of the vicariant range of Albinaria . The remaining species from Cyprus, Turkey and Greece constitute a well-supported monophyletic group. These two clades form geographical clusters, whereas Albinaria in the current definition (including ' A. '  hedenborgi ) has a disjunct distribution. ' A.' hedenborgi should therefore be classified with Cristataria , together with the morphologically similar and geographically close ' A. '  nadimi .  © 2005 The Linnean Society of London, Zoological Journal of the Linnean Society , 2005, 143 , 531−542.     

Intestinal Ultrastructure of Nerita picea (Mollusca: Gastropoda), an Intertidal Marine Snail of Hawaii

The neritid snail Nerita picea is a marine prosobranch mollusc which resides high in the intertidal zone on the Hawaiian Islands. Since other studies have shown considerable variations in molluscan gut histology and the relatively few recent ultrastructural reports have revealed novel cellular structures in the molluscan gastrointestinal tract, this investigation was directed toward ultrastructural clarification of the neritid intestine. Seven principal cell types constituted the intestinal architecture, including absorptive cells, zymogen cells, neural and endocrine cells, myocytes, pigment and gland cells. The intestinal epithelium was composed mainly of tall ciliated (9 plus 2 complement of microtubules) columnar absorptive cells which also possessed microvilli, extensive deposits of non-membrane-bound lipid-like droplets, and large reservoirs of glycogen-like granules. Less frequent, columnar zymogen cells contained numerous large zymogen secretory granules and possessed microvilli but not cilia. Small endocrine-like cells with secretory granules were observed basolaterally between some absorptive cells, resembling mammalian gut endocrine cells. Nerve fibers were prevalent in close association with the epithelial cells. A thin layer of non-striated muscle was present, as well as a serosally located gland composed of storage cells with a granular matrix and large granules.     

The Trochidae and Turbinidae of the Kermadec Ridge (Mollusca: Gastropoda)

Abstract

Taieria erebus (Gnaphosidae) was found to be a versatile predator: it captured insects both cursorially (away from webs) and kleptopar-asitically (on alien webs); it captured spiders in both the presence and absence of webs; and it also ate the eggs of host spiders (oophagy). When T. erebus invaded webs, it was as an aggressive mimic — it performed a repertoire of vibratory behaviours to lure the host spider. Although T. erebus pursued and captured spiders on diverse web-types, it was more effective as a predator when invading densely (rather than sparsely) woven cribellate and non-sticky webs, and was especially effective on non-cribellate sticky webs. Gnaphosids are traditionally referred to as hunting spiders, but T. erebus built a small prey-capture web. T. erebus also preyed on segestriid spiders, then used their webs to catch more prey, this being an unusual example of a spider using, as a tool for predation, the spinning-work of another species from an unrelated family. T. erebus used specialised behaviours to prey on nesting cursorial spiders. Prey was either grasped or stabbed; the venom of T. erebus was highly potent against spiders. Experiments indicated that vision was of little or no importance in the predatory behaviour of T. erebus. The behaviour of T. erebus is compared to that of Portia, a web-building salticid spider which is very versatile in its predatory behaviour and has acute vision. T. erebus is discussed in relation to hypotheses concerning gnaphosid and salticid evolution.     

Molecular phylogenetics of Caenogastropoda (Gastropoda: Mollusca)

Caenogastropoda is the dominant group of marine gastropods in terms of species numbers, diversity of habit and habitat and ecological importance. This paper reports the first comprehensive multi-gene phylogenetic study of the group. Data were collected from up to six genes comprising parts of 18S rRNA, 28S rRNA (five segments), 12S rRNA, cytochrome c oxidase subunit I, histone H3 and elongation factor 1alpha. The alignment has a combined length of 3995 base positions for 36 taxa, comprising 29 Caenogastropoda representing all of its major lineages and seven outgroups. Maximum parsimony, maximum likelihood and Bayesian analyses were conducted. The results generally support monophyly of Caenogastropoda and Hypsogastropoda (Caenogastropoda excepting Architaenioglossa, Cerithioidea and Campanilioidea). Within Hypsogastropoda, maximum likelihood and Bayesian analyses identified a near basal clade of nine or 10 families lacking an anterior inhalant siphon, and Cerithiopsidae s.l. (representing Triphoroidea), where the siphon is probably derived independently from other Hypsogastropoda. The asiphonate family Eatoniellidae was usually included in the clade but was removed in one Bayesian analysis. Of the two other studied families lacking a siphon, the limpet-shaped Calyptraeidae was associated with this group in some analyses, but the tent-shaped Xenophoridae was generally associated with the siphonate Strombidae. The other studied hypsogastropods with an anterior inhalant siphon include nine families, six of which are Neogastropoda, the only traditional caenogastropod group above the superfamily-level with strong morphological support. The hypotheses that Neogastropoda are monophyletic and that the group occupies a derived position within Hypsogastropoda are both contradicted, but weakly, by the molecular analyses. Despite the addition of large amounts of new molecular data, many caenogastropod lineages remain poorly resolved or unresolved in the present analyses, possibly due to a rapid radiation of the Hypsogastropoda following the Permian-Triassic extinction during the early Mesozoic.