Nematocyst sequestration within the family Fionidae (Gastropoda: Nudibranchia) considering ecological properties and evolution.

Aeolid nudibranchs are well-known for their ability to incorporate cnidarian nematocysts and use them for defense; this process is tightly linked with the feeding preferences of molluscs. As many nudibranch groups show signs of ecology-based adaptive radiation, studies of prey-based defensive mechanisms can provide valuable insight into details of nudibranch evolutionary history. The main goal of this study is to test the correlation of ecological traits, feeding mechanisms, and prey preferences with cnidosac fine morphology and to pinpoint the phylogenetic value of these traits. We study the cnidosac morphology in thirteen species-representatives of the main lineages within the family Fionidae s.l. The morphological analysis includes histological sections, transmission electron microscopy, confocal laser scanning microscopy, and scanning electron microscopy. For phylogenetic study, available molecular data from public repositories were used, and phylogenetic trees were produced based on Bayesian Inference and Maximum likelihood analysis for a concatenated dataset of three molecular markers (COI, 16S, H3). In general, fionid cnidosacs fit the common aeolid pattern, but among different species we detected a high variation in type of obtained nematocysts, their arrangement within cnidophages, and in number of cell types within cnidosacs. We report on presence of cellules speciale in the haemocoel of all studied species, and for the first time, we report on cells with chitinous spindles in the haemocoel of all fionids except Eubranchus. The function of both these cell types remains unknown. The loss of functional cnidosacs occurred at least three times within Fionidae, and in case of the genera Phestilla, Calma, and Fiona, this loss is linked to their non-cnidarian diet. The diversity of cnidosac fine structure within Fionidae s.l. correlates with that of the radular morphology and feeding preferences of each species. Prey shifts between cnidarian and non-cnidarian prey (both through evolutionary shifts and individual variation) rarely occur within Fionidae s.l.; however, microevolutionary shifts between different hydrozoan species within a single genus are more common. Cnidosac morphology demonstrates considerable resulting changes even when switching between similar hydrozoan species, or changing the feeding site on same prey species. These data indicate that cnidosac morphology likely follows microevolutionary prey shifts-in other words, it is affected by switches in prey species and changes in feeding sites with a single prey species. Thus, the cnidosac morphology may be a useful indicator when studying ecological features of particular species.

Transcriptomic sequence dataset of a potential new model species for studying biomineralization Onchidoris muricata (Nudibranchia, Gastropoda, Mollusca).

Onchidoris muricata is a widespread shell-less species of nudibranch molluscs, which has unique for Gastropoda skeletal elements - subepidermal calcite spicules. The general and fine morphology of the spicules, as well as their maturation process in ontogenesis, have been studied in detail by authors. The uniqueness of spicules lies in their intracellular formation and location under the ectodermal epithelium, which is more typical for deuterostomes. We present O. muricata as a potentially new model species for studying calcification of intracellular protein structure. A total of 96 individuals were collected in the Kandalaksha Bay of the White Sea, both manually and by scuba diving. All individuals were divided into three groups based on morphological characteristics such as specimens' size, spicule condition etc. This division suggests the existence of three stages in postembryonic ontogenesis of O. muricata reflecting the maturation of the spicule complex. Total RNA samples were isolated from three size groups of molluscs in three biological replicates. Libraries were prepared from the polyadenylated RNA fraction and sequenced at NovaSeq6000 (Illumina), yielding a total of 112.8 Gb of 150 bp paired-end reads, corresponding to almost 1,000-fold coverage of the transcriptome. Representative transcriptome assembled de novo with Trinity. In addition to obtaining the transcriptome sequences of O. muricata, differential expression analysis was also performed for these three size groups. This allows us to trace the dynamics of molecular and biological processes during the life of a mollusc. The obtained data can then be used as a reference transcriptome for closely related species, to study specific expressed genes, to identify various unique sequences, including protein-coding ones, to understand biological processes, including biomineralization and much more.

Ontogenetic dynamics of the nudibranch epithelium in Onchidoris muricata (O.F. Müller, 1776).

The integumentary system is the set of organs forming the outermost layer of an animal's body. It comprises the epithelium, muscles, and elements of connective and nerve tissue. The integument acts as a physical barrier between the external environment and the internal environment that serves to protect and maintain the body of the animal. The body of nudibranch mollusks undergo significant changes during ontogenesis, with the subepidermal space changing as the mollusk grows. As the extracellular subepidermal matrix is modified, the number of collagen fibers increases, muscles and nerves develop, and calcite spicules appear and grow. Yet, specific knowledge pertaining to the transformation of the epithelium is absent. In the present work, the ontogenetic dynamics of the surface epithelium of nudibranch mollusks are traced for the first time using Onchidoris muricata (O. F. Müller, 1776) during the postlarval stages of development. Ontogenetic changes in the epithelium of O. muricata were studied using a complex set of morphological methods. According to our data, the degree of modification to the epithelium in ontogenesis depends on individual body parts and is not consistent throughout. First x-cells were recognized as the probable precursors to sclerocytes.

Morphology of the buccal apparatus of Dendronotus frondosus (Gastropoda: Nudibranchia).

Nudibranchia is the order of Gastropoda, which is known for its high diversity of dietary specializations and a diversified morphology of the buccal armature. The characters of the buccal armature are adaptive to the feeding mechanisms and diet preferences, but they also are phylogenetically informative. The morphological differences in the buccal armature imply different mechanisms of the synthesis and morphology of the formation zones. Here, we report new data on the morphology of the buccal complex in a cladobranch species with broad radula, Dendronotus frondosus, using light microscopy, transmission and scanning elctron microscopy and computer-based three- dimensional reconstruction and compare them with those on other nudibranchs. We report the presence of protective chitinous spindles in the epithelia of the oral tube and esophagus, which suggests the presence of Cnidaria in the diet of the studied mollusc species. Similar to other Cladobranchia, the jaw plate of D. frondosus is synthesized by the layer of gnathoblasts located posteriorly in the epithelial fold. Near the zone of jaw synthesis, the gnathoblasts with concentric granular endoplasmic reticulum structures were found. This may indicate the turnover of gnathoblasts during the life of a specimen. Despite that D. frondosus has a broad radula, the zone of its synthesis does not split into two parts, unlike other molluscs with broad radulae. A single radular tooth is formed by a few cells, such as in other Nudibranchia. The number of odontoblasts per tooth in D. frondosus is not defined by a tooth size. At the same time, the linear dimensions of the cells secreting radular teeth depend on the size of the teeth synthesized by them.

Jaw morphology and function in Drilonereis cf. filum (Oenonidae, Annelida).

Representatives of the extant family Oenonidae (Annelida, Eunicida) have a prionognath jaw apparatus, with maxillae having forceps-like elements, a number of asymmetrical dentate plates and long slender carriers, which is characteristic of some fossil forms known from the Paleozoic epoch. Therefore, data on the fine structure and functional morphology of Oenonidae jaws are helpful for the interpretation of fossil materials. The fine structure of the jaw apparatus and the ventral pharyngeal organ is studied in one species of the Oenonidae (Annelida)-Drilonereis cf. filum. The material was collected in the soft bottom of Marseille Bay (Mediterranean) and examined with the help of TEM and histological techniques. A three-dimensional (3D) reconstruction was made from a complete series of semithin sections. The entire jaw apparatus is about 500 µm in length; it includes ventral mandibles and four pairs of maxillae, connected with long paired dorsal carriers and an unpaired ventral carrier. While retracted, it reaches the VIII-XI chaetigers. The most solid part of the maxillary apparatus, that is, maxillae I and II, are 2.5-5 µm thick. The plate consists of a monolithic array of merged scleroprotein granules in which perforations, that is, spaces remaining from microvilli, are visible; the basal part of the maxillary plate is a layer of loosely arranged collagen fibers penetrated with microvilli and has no signs of sclerotization. A study of the jaws of Drilonereis cf. filum showed the presence of common jaw patterns in Eunicida order. Like the jaws of Dorvilleidae, Eunicidae, Onuphidae, and Lumbrineridae, the jaws of Drilonereis are formed at the basis of a typical annelid cuticle's transformation with epi- and basicuticular layers, and its impregnation by merging scleroprotein granules. Through the nature of sclerotization, the jaws of D. cf. filum are similar to those of Dorvilleidae, Histriobdellidae, and the juvenile jaws of Mooreonuphis stigmatis (Onuphidae). Analysis of the 3D-reconstructions of the D. cf. filum jaw apparatus shows that the MxI of this species, and probably of other Oenonidae with dorsal and ventral carriers, can make grasping motions by fixing the joint of the right and left MxI in the two-door hinge type. In general, the overall structure of the jaw apparatus of D. cf. filum and the mechanics of its work shows greater similarity with that of Dorvilleidae than with the jaw apparatus of extant Labidognatha and Simmetrognatha (Onuphidae, Eunicidae, Lumbrineridae). The need for compactization of the jaw apparatus when moving in dense sediment or in the burrows is probably one of the factors determining its structure.

Main patterns of radula formation and ontogeny in Gastropoda.

Gastropoda is morphologically highly variable and broadly distributed group of mollusks. Due to the high morphological and functional diversity of the feeding apparatus gastropods follow a broad range of feeding strategies: from detritivory to highly specialized predation. The feeding apparatus includes the buccal armaments: jaw(s) and radula. The radula comprises a chitinous ribbon with teeth arranged in transverse and longitudinal rows. A unique characteristic of the radula is its continuous renewal during the entire life of a mollusk. The teeth and the membrane are continuously synthesized in the blind end of the radular sac and are shifted forward to the working zone, while the teeth harden and are mineralized on the way. Despite the similarity of the general mechanism of the radula formation in gastropods, some phylogenetically determined features can be identified in different phylogenetic lineages. These mainly concern shape, size, and number of the odontoblasts forming a single tooth. The radular morphology depends on the shape of the formation zone and the morphology of the subradular epithelium. The radula first appears at the pre- and posttorsional veliger stages as an invagination of the buccal epithelium of the larval anterior gut. The larval radular sac is lined with uniform undifferentiated cells. Each major phylogenetic lineage is characterized by a specific larval radula type. Thus, the docoglossan radula of Patellogastropoda is characterized by initially three and then five teeth in a transverse row. The larval rhipidoglossan radula has seven teeth in a row with differentiation into central, lateral, and marginal teeth and later is transformed into the adult radula morphology by the addition of lateral and especially marginal teeth. The taenioglossan radula of Caenogastropoda is nearly immediately formed in adult configuration with seven teeth in a row.

The rhipidoglossan radula: Radular morphology and formation in Nerita litterata Gmelin, 1791 (Neritimorpha, Neritidae).

The rhipidoglossan radula, consisting of numerous teeth in each transverse row, is characteristic of phylogenetically distant groups of gastropods, including Vetigastropoda, Neritimorpha and 'lower' Heterobranchia. Previous studies have revealed the main patterns in the formation of the rhipidoglossan radula of vetigastropods, the main feature of which is the division of the formation zone into two horns, where marginal teeth are formed by a multilayered epithelium (odontoblasts). This work is devoted to the study of the formation of the rhipidoglossan radula of Nerita litterata using light and electron microscopy. The data obtained show that, despite the different external morphology of the radular sac of neritids and vetigastropods, the radular sac of N. litterata, like that of vetigastropods, is divided into two parts, in which the marginal teeth are similarly formed by odontoblasts located in more than one layer. It seems probable that this complex, three-dimensional structure of the formation zone is associated with a broad radula with numerous elongate marginal teeth and could be characteristic of other gastropods with this type of radula. Additional supporting rods located along the odontoblasts and consisting of vacuolated cells were first discovered in Nerita.

Ontogenetic dynamics of the subepidermal spicule complex in Nudibranchia (Gastropoda): the case of Onchidoris muricata.

Spicules are mineral-based biocomposites skeletal structures that are widely distributed among phylogenetically distant groups of invertebrates (Porifera, Cnidaria, Mollusca, Echinodermata). Subepidermal spicules are formed under the ectodermal epithelium and are characterized for all groups except mollusks (Aplacophora, Polyplacophora, Bivalvia), their spicules are located on the surface of the body. However, one group of mollusks (Gastropoda: Heterobranchia) have unique subepidermal spicules that have never been detected above the ectodermal epithelium and similarly to those characterized for Porifera, Cnidaria and Echinodermata. Understanding subepidermal spicule formation in mollusks could help solve the question on the origin of spicules. Spicules in nudibranchs have been described for more than 150 years, yet ontogenetic dynamics of spicules have never been studied and the full mechanism of their formation remains unknown. Herein we investigate the spicule formation in different stages of postlarval development of the nudibranch Onchidoris muricata (O.F. Müller, 1776). For the first time, ontogenetic transformations of the spicule complex are described using experiments and different morphological methods. Our studies demonstrate that spicules of O. muricata form in the subepidermal space in early developmental stages immediately after veliger settlement. A single spicule forms inside a huge vacuole within a sclerocyte and remains there throughout the entire life of the specimen. Signs of spicule or sclerocyte migration under the epithelium in postlarval development was not found. Spicules only form during larval settlement, increasing only in size as development furthers. For the first time, spicule mineralization zones were detected at the tips of the spicules as well as the presence of collagen I in the overall composition of the spicules. Thus, our findings suggest that spicules form by an ectodermal cell that emerged under the ectodermal epithelium during the earliest stages of postlarval development.

The rhipidoglossan radula: Formation and development in Margarites helicinus Phipps, 1774 (Trochoidea, Vetigastropoda).

The gastropod radula exhibits exceptional morphological variability. Despite this enormous diversity, the main patterns of synthesis of the teeth and radula membrane, characteristic of different groups can be identified. The rhipidoglossan radula contains numerous teeth in each transverse row and was found in phylogenetically distant groups of Gastropoda. Studying radula formation through stages of ontogeny in species with this type of radula is important in determining the main patterns of radula formation in gastropods. We studied the formation of the radula during development of one species of trochid vetigastropod, Margarites helicinus, using light and electron microscopy as well as confocal laser scanning microscopy. The adult radula is formed in the blind end of the radular sac, which bifurcates into two horns in vetigastropods. The numerous marginal teeth are synthesized in these horns while the central and lateral teeth form in the region where the horns fuse. This morphology of the formation zone appears to be a common pattern for all vetigastropods. The juvenile radula of M. helicinus consists of seven teeth per transverse row and its formation in the radular sac differs significantly from that in the adult. In the juvenile, the formation zone of the radular sac is not split into two horns, and the teeth and radular membrane are synthesized by relatively few, uniform cells. This organization of the larval radular sac is thought to represent a widely occurring larval pattern potentially present in all groups of gastropods. It is associated with early formation of all organs by few cells rather than representing a phylogenetic trait.

The rhipidoglossan radula: Formation and morphology of the radula in Puncturella noachina (Linnaeus, 1771) (Fissurellidae, Vetigastropoda).

The rhipidoglossan radula, which is characterized by presence of a central tooth, several lateral teeth, and numerous (more than 10) long marginal teeth in each transverse row, is found in three different subclasses, that is, Vetigastropoda, Neritomorpha and "lower" Heterobranchia. Details of radula formation and its ultrastructure have not been studied in any species with a rhipidoglossan radula. For the first time, we present such data for one vetigastropod species, Puncturella noachina. The radula itself and the radula formation zone were studied using light and electron microscopy (scanning and transmission), as well as confocal laser scanning microscopy. We identify the major features of Vetigastropoda rhipidoglossan radula formation, that is: the posterior bifurcation of the radula formation zone, creating paired horns into which the zones of formation of the marginal teeth extend; the supporting structure in the radula formation zone extends ventrally to strengthen this division; the odontoblasts of the marginal teeth form a multi-layered epithelium; membranoblasts do not differ from odontoblasts in ultrastructure; in some membranoblasts and cells of the sub- and supraradular epithelium basal bodies were found in the apical regions of the cells.

Microbial associations of shallow-water Mediterranean marine cave Solenogastres (Mollusca).

The first cave-dwelling Solenogastres-marine shell-less worm-like mollusks-were sampled from Mediterranean marine caves floor silt in the Marseille area. The mollusks were 1.5 mm in length, had a transparent body with shiny spicules and appear to represent a new Tegulaherpia species. Electron microscopy revealed a high number of microbial cells, located on the surface of the spicules as well as in the cuticle of Tegulaherpia sp. The observed microbial cells varied in morphology and were unequally distributed through the cuticle, reaching a highest density on the dorsal and lateral sides and being practically absent on the ventral side. Next Generation Sequencing (NGS) of V4 region of 16S rRNA gene amplicons, obtained from the DNA samples of whole bodies of Tegulaherpia sp. revealed three dominating microorganisms, two of which were bacteria of Bacteroidetes and Nitrospirae phyla, while the third one represented archaea of Thaumarchaeota phylum. The Operational Taxonomic Unit (OTU), affiliated with Bacteroidetes was an uncultured bacteria of the family Saprospiraceae (93-95% of Bacteroidetes and 25-44% of the total community, depending on sample), OTU, affiliated with Nitrospirae belonged to the genus Nitrospira (8-30% of the community), while the thaumarchaeal OTU was classified as Candidatus Nitrosopumilus (11-15% of the community). Members of these three microbial taxa are known to form associations with various marine animals such as sponges or snails where they contribute to nitrogen metabolism or the decomposition of biopolymers. A similar role is assumed to be played by the microorganisms associated with Tegulaherpia sp.

New insights into the morphology and evolution of the ventral pharynx and jaws in Histriobdellidae (Eunicida, Annelida).

The jaw apparatus in several annelid families represents a powerful tool for systematic approaches and evolutionary investigations. Nevertheless, for several taxa, this character complex has scarcely been investigated, and complete comparative analyses of all annelid jaws are lacking. In our comprehensive study, we described the fine structure of the jaw apparatus and the ventral pharyngeal organ (VPO) in Histriobdella homari - a minute ectocommensal of lobsters putatively belonging to the Eunicida - using different comparative morphological approaches, including SEM, TEM, CLSM and subsequent 3D reconstruction. The H. homari jaw apparatus is composed of ventral paired mandibles and dorsal symmetrical maxillae consisting of numerous dental plates, ventral carriers and an unpaired dorsal rod, and the general assemblage and arrangement of the different parts are highly comparable to those of other eunicid families. The jaw ultrastructure of histriobdellids resembles that of the families Dorvilleidae and (juvenile) Onuphidae. Furthermore, our data reveal that in the process of development of the jaw apparatus, the mandibles, maxillae II and unpaired dorsal rod are formed first, and the remaining maxillae and ventral carriers appear later. Notably, the muscular apparatus differs from that in Dorvilleidae and Onuphidae in terms of the number and arrangement of muscle fibers encompassing the jaws - not only because of the very small size of Histriobdella but also because histriobdellid maxillary protraction occurs due to straightening of the dorsal rod and thus requires a different muscular scaffold. Based on our investigations, the arrangement of the muscular apparatus of the jaws, the presence of paired ventral carriers and the dorsal rod, and the morphology of the ventral pharyngeal organ represent a histriobdellid autapomorphy. Our datasets form a basis for further comparative analyses to elucidate the evolution of Eunicida and jaw-bearing Annelida.

Functional morphology and post-larval development of the buccal complex in Eubranchus rupium (Nudibranchia: Aeolidida: Fionidae).

Nudibranch molluscs represent an interesting model group to study the evolution of feeding apparatus and feeding modes, being characterized by specialized buccal complex in combination with extremely diverse dietary preferences and multiply prey shifts in evolutionary history. However, the plasticity of the buccal complex morphology in response to diet and specific feeding modes remains understudied. Here we study the general morphology and ontogenesis of the buccal complex in Eubranchus rupium (Nudibranchia: Fionidae). Specific goals are to provide a detailed description of buccal structures morphology in post-larval stages, suggest the feeding mechanism and discuss the phylogenetic value of the morphological characteristics of buccal armature within the genus Eubranchus. Methods included in vivo observations of the feeding process for E. rupium, light microscopic methods, scanning electron microscopy (SEM), transmission electron microscopy (TEM) and confocal laser scanning microscopy (cLSM). According to our results, E. rupium is a mechanical driller, boring holes in hydrozoan perisarc and sucking internal content. The mechanical drilling is supplied by functionally uniserial radula with plate-like laterals teeth of exclusively supportive function and by massive buccal musculature. Comparative phylogeny-based analysis suggests that the drilling feeding mode is common for the genus Eubranchus and indicates radular characters may have a high phylogenetic signal. The buccal complex morphology and feeding mode were found to be similar in both adults and post-metamorphic specimens, its general structures occur even in settled veligers. Juveniles and adults compete for food source, but the competition is smoothed due to characteristics of prey species growth and life cycle.

Radula formation in two species of Conoidea (Gastropoda).

The radula is the basic feeding structure in gastropod molluscs and exhibits great morphological diversity that reflects the exceptional anatomical and ecological diversity occurring in these animals. This uniquely molluscan structure is formed in the blind end of the radular sac by specialized cells (membranoblasts and odontoblasts). Secretion type, and the number and shape of the odontoblasts that form each tooth characterize the mode of radula formation. These characteristics vary in different groups of gastropods. Elucidation of this diversity is key to identifying the main patterns of radula formation in Gastropoda. Of particular interest would be a phylogenetically closely related group that is characterized by high variability of the radula. One such group is the large monophyletic superfamily Conoidea, the radula of which is highly variable and may consist of the radular membrane with five teeth per row, or the radular membrane with only two or three teeth per row, or even just two harpoon-like teeth per row without a radular membrane. We studied the radulae of two species of Conoidea (Clavus maestratii Kilburn, Fedosov & Kantor, 2014 [Drilliidae] and, Lophiotoma acuta (Perry, 1811) [Turridae]) using light and electron microscopy. Based on these data and previous studies, we identify the general patterns of the radula formation for all Conoidea: the dorsolateral position of two groups of odontoblasts, uniform size, and shape of odontoblasts, folding of the radula in the radular sac regardless of the radula configuration. The morphology of the subradular epithelium is most likely adaptive to the radula type.

General morphology and ultrastructure of the radula of Testudinalia testudinalis (O. F. Müller, 1776) (Patellogastropoda, Gastropoda).

The radular morphology of the patellid species Testudinalia testudinalis (O. F. Müller, 1776) from the White Sea was studied using light, electron, and confocal microscopy. The radula is of the docoglossan type with four teeth per row and consisting of six zones. We characterize teeth formation in T. testidinalis as follows: one tooth is formed by numerous and extremely narrow odontoblasts through apocrine secretion; this initially formed tooth consists of numerous vesicles; the synthetic apparatus of the odontoblasts is localized in the apical and central parts of the cells throughout the cytoplasm and is penetrated by microtubules which are involved in the transport of the synthesized products to the apical part of the odontoblast; the newly formed teeth consist of unpolymerized chitin. Mitotic activity is located in the lateral parts of the formation zone. The first four rows contain an irregular arrangement of teeth, but the radular teeth are regularly arranged after the fifth row. The irregularly arranged teeth early on could be a consequence of the asynchronous formation of teeth and the distance between the odontoblasts and the membranoblasts. The morphological data obtained significantly expands our knowledge of the morphological diversity of the radula formation in Gastropoda.

Drilling in the dorid species Vayssierea cf. elegans (Gastropoda: Nudibranchia): Functional and comparative morphological aspects.

The drilling mode of feeding is known from two clades of Gastropoda: Caenogastropoda and Heterobranchia. However, the level of convergence and parallelism or homology among these two lineages is unclear. The morphology of the buccal complex is well studied for drilling caenogastropods, but poorly known for drilling nudibranchs. It is also unclear whether the drilling feeding mechanism is similar between inside gastropods. Accordingly, a comparison between the feeding mechanisms of drilling nudibranchs and caenogastropods can help to understand the evolutional trends inside gastropods. In this study, we redescribe the morphology of the buccal complex of drilling dorid nudibranch Vayssierea cf. elegans, and compare it to that of previous investigations on this species and closely related dorid species. We describe the feeding mechanism of this species based on the obtained morphological and literature data and compare it to the feeding mechanisms described for drilling caenogastropods. The feeding apparatus of Vayssierea cf. elegans corresponds to the general morphology of the dorid buccal complex; that is, it has a similar arrangement of the buccal musculature and pattern of radular morphology. However, there are also adaptations to the drilling feeding mode similar to those found in Caenogastropoda: that is, specialized dissolving glands and lateral teeth with elongated pointed cusps; and even Sacoglossa: the specialized muscle for sucking. The feeding process of Vayssierea cf. elegans includes the same two stages as those described for drilling caenogastropods: (a) the boring stage, which is provided by mechanical and chemical activity, and (b) the swallowing stage.

Fine morphology of the jaw apparatus of Puncturella noachina (Fissurellidae, Vetigastropoda).

Jaws of various kinds occur in virtually all groups of Mollusca, except for Polyplacophora and Bivalvia. Molluscan jaws are formed by the buccal epithelium and either constitute a single plate, a paired formation or a serial structure. Buccal ectodermal structures in gastropods are rather different. They can be nonrenewable or having final growth, like the hooks in Clione (Gastropoda, Gymnosomata). In this case, they are formed by a single cell. Conversely, they can be renewable during the entire life span and in this case they are formed by a set of cells, like the formation of the radula. The fine structure of the jaws was studied in the gastropod Puncturella noachina. The jaw is situated in the buccal cavity and consists of paired elongated cuticular plates. On the anterior edge of each cuticular plate there are numerous longitudinally oriented rodlets disposed over the entire jaw surface and immersed into a cuticular matrix. The jaw can be divided into four zones situated successively toward the anterior edge: 1) the posterior area: the zone of formation of the thick cuticle covering the entire jaw and forming the electron-dense outer layer of the jaw plate; 2) the zone of rodlet formation; 3) the zone of rodlet arrangement; and 4) the anterior zone: the free scraping edge of the plate, or the erosion zone. In the general pattern of jaw formation, Puncturella noachina resembles Testudinalia tessulata (Patellogastropoda) studied previously. The basis of the jaw is a cuticular plate formed by the activity of the strongly developed microvillar apparatus of the gnathoepithelium. However, the mechanism of renewal of the jaw anterior part in P. noachina is much more complex as its scraping edge consists not just of a thick cuticular matrix rather than of a system of denticles being the projecting endings of rodlets.