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.

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.

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.

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.

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.