The phylogenetic position of the heteropod Atlanta gaudichaudi Souleyet (Mollusca, Gastropoda), a spermatological investigation

The male adult, the spermatophore and the spermatozoa of Atlanta gaudichaudi are described. No parasperm are demonstrable. The ultrastructure of the spermatozoa is shown to differ fundamentally from that of the Archaeogastropoda and Neritopsina and to conform broadly with that of the Mesogastropoda (excepting Cyclophoracea, Viviparacea and Cerithacea), the Neogastropoda and the epitonacean heterogastropods. In having a nuclear fossa, containing the axoneme, which penetrates the entire length of the nucleus as the intranuclear canal, the spermatozoon especially resembles that of some members of the Rissoacea, Littorinacea, Eulimacea, Tonnacea, Volutacea and Conacea. Absence, in Atlanta of type 7 parasperm ( sensu Koike 1985), which characterize the Calyptraeacea, Cypraeacea, Tonnacea, Buccinacea, Volutacea (including the Mitracea) and most Conacea, or of type 8 parasperm of the Conidae, contraindicates heteropod relationship with these taxa. Relationship to the Rissoacea cannot be ruled out, though presence of oligopyrene, cusperm-like sperm in these (Bythiniidae) and pterotracheid heteropods is a questionable indication of affinity. Relationship of heteropods with the Littorinacea is suggested by similarity of the sperm, presence in carinariid heteropods and littorinids of nurse cells, and apparent homology of the male reproductive apparatus in both groups. The latter two similarities require confirmation, however. It is suggested that deposition of spermatophores on the exterior of the female in atlantid heteropods is a primitive mode of sperm transfer compatible with relationship with the Littorinacea.     

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.     

Phylogeny of the Cocculinoidea (Mollusca, Gastropoda)

Abstract. The superfamily Cocculinoidea is a group of marine, deep-water, limpet-like gastropods. Recent speculation surrounding their affinities has concentrated on their placement within the Gastropoda. However, phylogenetic relationships within the Cocculinoidea, especially the monophyly of families and genera within the group, remain poorly understood. Phylogenetic analysis of 31 morphological characters for 15 cocculinoidean taxa and 2 outgroups resulted in a single most parsimonious tree, length=70, CI=0.62, and RI=0.71. Monophyly of the Cocculinoidea, Cocculinidae, and the genera Cocculina and Coccopigya was supported; Paracocculina and Coccocrater were found to be paraphyletic. Character optimization demonstrates that many characters often cited as diagnostic of various taxa, are often homoplastic and/or synapomorphies at different hierarchical levels.     

Heterobranchia (Gastropoda) from the Jurassic Deposits of Russia

Small gastropods from the Jurassic deposits of the European Russia, united into the lower Heterobranchia or Allogastropoda, are described. The families of Ampezzanildidae, Cimidae, Cornirostridae, Ebalidae, and Stuoraxidae are distinguished, and a set of taxa is given without reference to a definite family. The family Ampezzanildidae from the Jurassic deposits is described for the first time on the basis of the mass material, ascribed to the new genus Zizipupa gen. nov. with the sole species of Z. costata sp. nov. The family Cimidae includes the genera of Cristalloella, Rotfanella, Urlocella, and Unzhispira gen. nov. with species C. spiralocostata (Gründel, 1998), R. gerasimovi sp. nov., R. reticulata sp. nov., Urlocella undulata sp. nov., and Unzhispira minuta sp. nov. The genus Heteronatica gen. nov. is included into the family Cornirostridae. This genus is the first siphonostomatous representative of the given family including the sole long-living species H. globosa sp. nov., which is subdivided into subspecies H. globosa globosa and H. globosapromota subsp. nov. The family Ebalidae is represented by the genus Ebala, shells of which are distributed from the Middle Oxfordian to the Middle Volgian. The family Stuoraxidae is described based on two genera Stuoraxis and Aneudaronia gen. nov., including species of S. crassa sp. nov. and A. elegans sp. nov. The genus Doggerostra, which fits into different families of Heterobranchia in terms of shell morphology is represented by the species D. riedeli Gründel, 1998, which was previously known from the Bathonian and Callowian deposits in Poland and Germany, as well as form the Upper Jurassic interval in the Russian Plate. The Middle Volgian subspecies D. riedeli affinis subsp. nov. is distinguished in the composition of this species. In addition, the genus Masaevia gen. nov. with the sole species of M. sinistra sp. nov. is described. Due to so specific shell morphology of this genus, its position in the Heterobranchia system is unclear. It is not improbable that we deal with small planktonic gastropods of protoconchs of unknown group of planktonic or benthic gastropods.     

The complete mitochondrial genome of the nudibranch Roboastra europaea (Mollusca: Gastropoda) supports the monophyly of opisthobranchs

The complete nucleotide sequence (14,472 bp) of the mitochondrial genome of the nudibranch Roboastra europaea (Gastropoda: Opisthobranchia) was determined. This highly compact mitochondrial genome is nearly identical in gene organization to that found in opisthobranchs and pulmonates (Euthyneura) but not to that in prosobranchs (a paraphyletic group including the most basal lineages of gastropods). The newly determined mitochondrial genome differs only in the relative position of the trnC gene when compared with the mitochondrial genome of Pupa strigosa, the only opisthobranch mitochondrial genome sequenced so far. Pupa and Roboastra represent the most basal and derived lineages of opisthobranchs, respectively, and their mitochondrial genomes are more similar in sequence when compared with those of pulmonates. All phylogenetic analyses (maximum parsimony, minimum evolution, maximum likelihood, and Bayesian) based on the deduced amino acid sequences of all mitochondrial protein-coding genes supported the monophyly of opisthobranchs. These results are in agreement with the classical view that recognizes Opisthobranchia as a natural group and contradict recent phylogenetic studies of the group based on shorter sequence data sets. The monophyly of opisthobranchs was further confirmed when a fragment of 2,500 nucleotides including the mitochondrial cox1, rrnL, nad6, and nad5 genes was analyzed in several species representing five different orders of opisthobranchs with all common methods of phylogenetic inference. Within opisthobranchs, the polyphyly of cephalaspideans and the monophyly of nudibranchs were recovered. The evolution of mitochondrial tRNA rearrangements was analyzed using the cox1+rrnL+nad6+nad5 gene phylogeny. The relative position of the trnP gene between the trnA and nad6 genes was found to be a synapomorphy of opisthobranchs that supports their monophyly.     

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.     

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.     

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.     

Linnaeus' Neptunea (Mollusca: Gastropoda)

Three species of the arcto-boreal, large gastropod Neptunea , described by Linnaeus in 1758 and 1771, occur in large numbers over wide areas of the inshore North Atlantic and adjacent Arctic seas and are conspicuous among Pliocene and Pleistocene molluscs in the Icelandic, North Sea, and western Mediterranean basins. Selections of lectotypes for these species from shells in the collection of the Linnean Society of London, and designations of their type localities, establish the identity of Linnaeus' neptunes and more accurately determine their geographic and geologic distribution. The geographic range of Neptunea (Neptunea) antiqua (L.), the type species, now extends from southern Norway to the northern Biscay coast of France and from the westernmost Baltic Sea to southwestern Ireland; this species also occurs in Pliocene-Holocene marine deposits in West and East Germany, Sweden, the Netherlands, England and France. Its type locality is determined to be the North Sea. N. (Neptunea) despecta (L.) lives in the eastern Canadian Arctic, off southern Greenland, the Barents Sea, and North Atlantic as far south as Massachusetts and Portugal; it also occurs in Pliocene-Holocene strata of eastern Canada, east-central Greenland, Norway (including Svalbard), the Soviet Union, Sweden and England. Its type locality is determined to be the postglacial deposits at Uddevalla in southwestern Sweden. N. (Sulcosipho) contraria (L.) now extends from the southern Biscay coast of France to Cape Spartel, Morocco; this species also occurs in Pleistocene and lower Holocene sequences of the western Mediterranean. Its type locality is determined to be Vigo Bay, Spain. A closely related fossil species, N. (S.) angulata (S. V. Wood), occurs in Pliocene and Pleistocene deposits of the North Sea basin.     

Molecular confirmation of the occurrence of Elysia cf. tomentosa (Mollusca: Heterobranchia) in the Persian Gulf

The molluscan fauna of the Persian Gulf has recently been relatively well documented, yet there are few records of heterobranch sea slugs (opisthobranchs) from the Arabian parts and no report from the Iranian waters. Here we report for the first time the occurrence of one of these molluscs in the northern Persian Gulf (Bandar Abbas, Iran). Sacoglossan specimens were collected in association with the seaweed, Caulerpa sertularioides. Since morphological attributes were not adequately reliable for species identification, molecular approaches were carried out. Maximum-likelihood and Bayesian Inference analysis of partial DNA sequences of the mitochondrial cytochrome c oxidase subunit I (COI) locus were used for DNA barcoding of large-bodied specimens of Elysia. All Persian Gulf specimens were genetically confirmed as Elysia cf. tomentosa sp. 5, one of at least five morphologically similar but genetically distinct species in the taxonomically challenging and unresolved E. tomentosa complex. This species has previously been recorded only from Australia and Thailand and our finding adds another distant point to the geographic distribution of this species.     

The taxonomy of two host-specific, cryptic dendronotoid nudibranch species (Mollusca:Gastropoda) from Australia including a new species description

Two host-specific, extremely cryptic, small dendronotoid nudibranchs from Australia are described and classified, and their phylogenetic position within the superfamily is discussed- Lomanotus vermiformis Eliot, 1908 subsists exclusively on the hydrozoan Lytocarpus philippinus (Kirchenpauer, 1872). Lomanotus stauberi Clark & Goetzfried, 1976 is newly synonymized with L. vermiformis . Marioniopsis platyctenea sp. nov. subsists exclusively on the alcyonacean Parerythropodium hicksoni Utinomi, 1972. Marianina rosea (Pruvot-Fol, 1930) is relocated to the Tritoniidae (wherein it forms a monotypic subfamily, Marianinae) because of possession of palmate rhinophoral clavi, the principal apomorphy of the Tritoniidae. The validity of using the form of the digestive gland as a basis for dividing the Tritoniidae is questioned since it is acknowledged that this organ has progressed from a solid (holohepatic) arrangement to a dispersed (cladohepatic) arrangement in parallel in several major nudibranch lineages; form thus offers merely homoeoplaseous phylogenetic characters. Separation of right and left digestive glands appears to be one of the first steps in the simultaneous and interdependent evolutionary processes of internal detorsion and external 'aeolidization' in nudibranchs.     

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.     

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.     

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.     

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.     

Nuclear morphogenesis during spermiogenesis in the nudibranch molluscHypselodoris tricolor (Gastropoda,opisthobranchia)

An electron microscope study was carried out on Hypselodoris tricolor spermatids to describe the development of the nuclear morphogenesis and investigate the possible cause(s) of the change in the shape of the spermatid nucleus during spermiogenesis. Three different stages may be distinguished in the course of the nuclear morphogenesis on the basis of the morphology and inner organization of the nucleus. Stage 1 spermatid nuclei are spherical or ovoid in shape and the nucleoplasm finely granular in appearance. Stage 2 nuclei exhibit a disc- or cup-shaped morphology, and the chromatin forms short, thin filaments. During stage 3, a progressive nuclear elongation takes place, accompanied by chromatin rearrangement, first into fibers and then into lamellae, both formations helically oriented. A row of microtubules attached to the nuclear envelope completely surrounds the nucleus. Interestingly, the microtubules always lie parallel to the chromatin fibers adjacent to them. Late stage 3 spermatids show the highest degree of chromatin condensation and lack the manchette at the end of spermiogenesis. Our findings indicate the existence of a clear influence exerted on the chromatin by the manchette microtubules, which appear to be involved in determining the specific pattern of chromatin condensation in Hypselodoris tricolor.     

Novel copulatory structures and reproductive functions in Amphiboloidea (Gastropoda, Heterobranchia, Pulmonata)

Abstract. Two new families of amphiboloid gastropods were recently erected, largely due to the very distinct structural and functional configurations of their distal reproductive systems. The spermovipositors in monaulic Salinator spp. (Amphibolidae) are functionally unique in Gastropoda, and it is suggested that they perform both oviposition and sperm transfer/receipt during non-reciprocal copulation. In contrast, the remarkably complex penises in syntremous diaulic Phallomedusa spp. (Phallomedusidae) are used exclusively for sperm transfer during reciprocal copulation. The phallomedusid penis is structurally unique among gastropods because it is protruded by pivoting around a fixed point rather than extending by eversion, and bears elaborate appendages to improve traction during copulation. The complex and functionally ambiguous distal reproductive structures of Maningrida (Maningrididae) differ from other amphibolids in the arrangement and structure of their prostate and copulatory structures. Genital armature of varying complexity occurs in all amphiboloidean families. There may be a correlation between the absence of a long-term sperm storage structure (spermatheca) and the evolutionary diversification of copulatory structures in these gastropods.     

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 .