Discovery and characterization of a transient chaetal gland during the development of Capitella teleta (Sedentaria: Annelida).

Chaetae are among the most extensively studied structures in polychaetes, serving as a defining morphological trait for annelids. Capitella teleta stands out as one of the few established annelid models for developmental and morphological studies, thus receiving significant scholarly attention. In this study, we unveil a previously unnoticed glandular structure associated with chaetae within the larvae of C. teleta. Our investigations demonstrate the absence of comparable structures in the chaetal follicles of adults and juveniles (older than 1 week), as well as during active chaetogenesis, underscoring the transient nature of these glands. This indicates that larval chaetal follicles transform into a gland that later disappears. Utilizing histology and transmission electron microscopy, we characterized these glands. Our findings underscore the diversity of chaetal ultrastructure in annelids and show that, even in well-studied species, novel morphological details can be found. We emphasize the importance of examining various life-history stages to capture such transient morphological features. This work lays a crucial morphological foundation and deepens our understanding of chaetae and chaetogenesis in C. teleta, paving the way for more accurate interpretations of future experimental studies on chaetogenesis in this species.

Morphological and molecular features of early regeneration in the marine annelid Ophryotrocha xiamen.

Regeneration capability varies in the phylum Annelida making them an excellent group to investigate the differences between closely related organisms. Several studies have described the process of regeneration, while the underlying molecular mechanism remains unclear, especially during the early stage (wound healing and blastema formation). In this study, the newly identified Ophryotrocha xiamen was used to explore the early regeneration. The detailed morphological and molecular analyses positioned O. xiamen within 'labronica' clade. We analyzed the morphological changes during regeneration process (0-3 days post amputation) and molecular changes during the early regeneration stage (1 day post amputation). Wound healing was achieved within one day and a blastema formed one day later. A total of 243 DEGs were mainly involved in metabolism and signal transduction. Currently known regeneration-related genes were identified in O. xiamen which could help with exploring the functions of genes involved in regeneration processes. According to their conserved motif, we identified 8 different Hox gene fragments and Hox5 and Lox2 were found to be absent in early regeneration and during regular growth. Our data can promote further use of O. xiamen which can be used as an experimental model for resolving crucial problems of developmental biology in marine invertebrates.

All for one: changes in mitochondrial morphology and activity during syncytial oogenesis†.

The syncytial groups of germ cells (germ-line cysts) forming in ovaries of clitellate annelids are an attractive model to study mitochondrial stage-specific changes. Using transmission electron microscopy, serial block-face scanning electron microscopy, and fluorescent microscopy, we analyzed the mitochondria distribution and morphology and the state of membrane potential in female cysts in Enchytraeus albidus. We visualized in 3D at the ultrastructural level mitochondria in cysts at successive stages: 2-celled, 4-celled, 16-celled cysts, and cyst in advanced oogenesis. We found that mitochondria form extensive aggregates-they are fused and connected into large and branched mitochondrial networks. The most extensive networks are formed with up to 10 000 fused mitochondria, whereas individual organelles represent up to 2% of the total mitochondrial volume. We classify such a morphology of mitochondria as a dynamic hyperfusion state and suggest that this can maintain their high activity and intensify the process of cellular respiration within the syncytial cysts. We found some individual mitochondria undergoing degradation, which implies that damaged mitochondria are removed from networks for their final elimination. As growing oocytes were shown to possess less active mitochondria than the nurse cells, the high activity of mitochondria in the nurse cells and their dynamic hyperfusion state are attributed to serve the needs of the growing oocyte. In addition, we measured by calorimetry the total antioxidant capacity of germ-line cysts in comparison with somatic tissue, and it suggests that antioxidative defense systems, together with mitochondrial networks, can effectively protect germ-line mitochondria from damage.

New data on echiuran anatomy and histology: the case of Lissomyema mellita (Annelida: Thalassematidae).

Echiura is small group of unsegmented marine worms that are sometimes abundant in the benthos of all areas of the World Ocean. The study of echiuran morphology and anatomy is important for understanding echiuran biology and the function of benthic communities where echiurans dominate. The current study used paraffin histology, scanning electron microscopy, and 3D reconstruction to describe the anatomy of organ systems in Lissomyema mellita, which is within the tribe Thalassematini of the subfamily Thalassematinae. The body of L. mellita consists of a trunk and a proboscis. The trunk terminates at the anal lobe, which contains many large glands, the secretions of which may be used for processing of the hard substratum where studied animals were found. The proboscis has lateral ciliated grooves that are < 7 µm wide and are probably used for collecting and separating food particles. The coelom is divided into the coelom of the proboscis and the coelom of trunk. The location of proboscis coelom may suggest that the proboscis is derived from the oral segment of the body of a hypothetical segmented echiuran ancestor. The shortness and histological uniformity of the foregut indicates that L. mellita feeds on soft food particles that do not require mechanical processing. The circulatory system has a complex organization and contains several vascular elements that have not been previously described in echiurans: voluminous plexus around the foregut, voluminous dorsal and ventral lacunes, and additional vessels between neurointestinal and circular blood vessels. The dorsal blood vessel is very short; the circumchaetael vessel is absent. Most of the anatomical peculiarities of L. mellita appear to be related to its biology. The newness of the data on the organization of the circulatory system reflects the general scarcity of knowledge on echiuran anatomy. The newly discovered features should be incorporated into general schemes of echiuran organization.

Unravelling the ultrastructure and mineralogical composition of fireworm stinging bristles.

Amphinomid fireworms are notorious for their stinging dorsal bristles (notochaetae), but it is still unclear whether the irritation they cause is merely mechanical or if the notochaetae contain toxins. Furthermore, although fireworm chaetae have always been described as calcareous, their composition has never been investigated to date and strong debates are ongoing on their internal structure. Unravelling the native ultrastructure and composition of fireworm chaetae is the first crucial step to assess whether the hypothesis of toxin vehiculation could be fully considered. We examined for the first time the chemical and mineralogical composition, the ultrastructure and the external structure of the dorsal and ventral chaetae of the large species Hermodice carunculata. All the measurements were carried out on samples prepared without the use of chemical reagents, except for those targeted to investigate if decalcification altered the ultrastructure of the chaetae. A crystal-chemical strategy, combining chemical, diffraction and thermal analyses clearly showed the occurrence of crystalline calcium carbonate and clusters of phosphatic amorphous material. Scanning electron micrographs and energy dispersive X-ray measurements showed that the dorsal chaetae have an extremely shallow insertion point in the body respect to the ventral chaetae, that could facilitate the release of the notochaetae in the environment. Their proximal part is characterized by canals with a hexagonal pattern rich in Ca and P, followed by a large cavity upwards. The harpoon-shaped ends and the central canals of the notochaetae completely disappeared after exposure to EDTA. The notochaetae are hollow and may be able to vehicle toxins. The absence of the honeycomb pattern in the distal part of the notochaetae and their slenderness probably contribute to their brittleness and high sensitivity to breakage on contact. These observations constitute keystone understandings to shed light on fireworm defensive and offensive capacities and their ecological success.

On the genus Sabidius Strelzov, 1973 (Annelida: Paraonidae), with a redescription of the type species and the description of a new species.

Two species of Sabidius Strelzov, 1973 were collected during a benthic survey, from 10-3,000 m deep, in Espírito Santo Basin, off southeastern Brazilian coast. Those species are S. cornatus (Hartman, 1965) and S. antennatus sp. nov. The genus Sabidius was monotypic until the present study, in which a new species is proposed, together with a redescription of the type species, with expansion of the geographic distribution of the genus and of the type species. The diagnostic feature of the genus is the morphology of prostomium, with crested anterior margin. The main feature that differentiates the two species within the genus is the presence/absence of a median antenna, which is present in the new species described in the present study and absent in Sabidius cornatus.

Description of ovary organization and oogenesis in a phreodrilid clitellate.

Phreodrilidae is a small family uniting about 50 species of minute freshwater clitellate annelids inhabiting mainly the Southern hemisphere. Other than the male and spermathecal genitalia, their internal organization is poorly known. Here, we present results of our study of the ovaries and oogenesis in Insulodrilus bifidus, a phreodrilid from Western Australia using light and electron microscopy. The ovaries are paired and located in segment XII. They are inconspicuous and composed of several (10-12) spherical germ-line cysts loosely interconnected by flattened somatic cells. The cysts usually comprise 32 germ cells and each cell is connected via a cytoplasmic bridge (ring canal) to the central cytoplasmic mass (the cytophore). In ovaries, germ cells in a given cyst develop in full synchrony. However, there is no synchrony among cysts, so there is a developmental gradient of cysts (from oogonial to early meiotic) along the longitudinal ovary axis. Within the cysts that are located in the distal end of the ovary the synchrony is finally lost and interconnected cells diversify into two morphologically distinct categories: an oocyte and 31 nurse cells. Such cysts detach from the ovaries and further development occurs within the body cavity. The oocyte gathers nutrients, mainly in form of yolk spheres, whereas nurse cells grow slightly and do not gather yolk. Organelles such as ribosomes, mitochondria and endoplasmic reticulum pass freely through the ring canals and are present within the cytophore, which suggests cytoplasmic transfer towards the oocyte. The formation of female germ-line cysts equipped with cytophore and cells differentiated into oocyte and nurse cells matches the general pattern of oogenesis found in clitellates. In details, the ovary organization and oogenesis found in I. bifidus resembles the situation described in some representatives of Naidinae and Enchytraeidae.

Protonephridial Excretory System in Vestimentifera (Siboglinidae, Annelida).

Ultrastructural study of the excretory tree of vestimentifera Ridgeia piscesae has shown that it consists of tubules that are blind at their distal ends. The tubules are lined with ciliated cells and have one or two multiciliated terminal cell(s) at the distal ends. In the tubule walls, there are putative ultrafiltration sites. The excretory tree tubules are interpreted as the secondary protonephridia.

Micromorphology of ovaries and oogenesis in Grania postclitellochaeta (Clitellata: Enchytraeidae).

The genus Grania comprises over 70 species of exclusively marine clitellate annelids belonging to the family Enchytraeidae. Morphologically, this genus is well separated from other enchytraeids, with thick cuticles, anterior segments I-IV fused into a "head", chaetal bundles consisting only of one stout chaeta, and reduction of circular musculature. The aim of the present study is to describe the ovary organization and the course of oogenesis in Grania postclitellochaeta, and to compare it with other known systems of ovary organization and oogenesis in clitellate annelids, especially in enchytraeids. Generally, oogenesis in G. postclitellochaeta can be divided into two phases: (i) early stages of oogenesis, occurring within the paired ovaries - each ovary is similar to a bunch of grapes, where each 'lobe' is a germ-line cyst enveloped by flat somatic cells, and (ii) oogenesis proper, which takes place within the body lumen where each growing oocyte is accompanied by its own group of nurse cells. Germ cells are interconnected by cytoplasmic channels (intercellular bridges, ring canals) and form syncytial cysts. As in other clitellate annelids, the cyst center contains a common cytoplasm (cytophore) to which each cell is connected by one ring canal only. Initially, within the ovary, all interconnected cells develop synchronously and are morphologically similar. At the time when the cysts detach from the ovary, one of the interconnected cells begins to gather nutrients, grows and becomes an oocyte, whereas the rest of the cells (nurse cells) do not continue meiosis and instead seem to provide the oocyte with macromolecules and cell organelles. Analysis of serial sections reveals that cysts are always composed of 16 cells - one oocyte and fifteen nurse cells. A comparative analysis showed that almost all features of oogenesis in G. postclitellochaeta are similar to that in other representatives of Enchytraeidae (mainly Enchytraeus albidus), suggesting evolutionary conservation of the process across this family.

Ammonia excretion in the marine polychaete Eurythoe complanata (Annelida).

Ammonia is a toxic waste product from protein metabolism and needs to be either converted into less toxic molecules or, in the case of fish and aquatic invertebrates, excreted directly as is. In contrast to fish, very little is known regarding the ammonia excretion mechanism and the participating excretory organs in marine invertebrates. In the current study, ammonia excretion in the marine burrowing polychaete Eurythoe complanata was investigated. As a potential site for excretion, the 100-200 µm long, 30-50 µm wide and up to 25 µm thick dentrically branched, well ventilated and vascularized branchiae (gills) were identified. In comparison to the main body, the branchiae showed considerably higher mRNA expression levels of Na+/K+-ATPase, V-type H+-ATPase, cytoplasmic carbonic anhydrase (CA-2), a Rhesus-like protein, and three different ammonia transporters (AMTs). Experiments on the intact organism revealed that ammonia excretion did not occur via apical ammonia trapping, but was regulated by a basolateral localized V-type H+-ATPase, carbonic anhydrase and intracellular cAMP levels. Interestingly, the V-type H+-ATPase seems to play a role in ammonia retention. A 1 week exposure to 1 mmol l-1 NH4Cl (HEA) did not cause a change in ammonia excretion rates, while the three branchial expressed AMTs showed a tendency to be down-regulated. This indicates a shift of function in the branchial ammonia excretion processes under these conditions.

Cosmopolitan or Cryptic Species? A Case Study of Capitella teleta (Annelida: Capitellidae).

Capitella teleta Blake et al., 2009 is an opportunistic capitellid originally described from Massachusetts (USA), but also reported from the Mediterranean, NW Atlantic, and North Pacific, including Japan. This putatively wide distribution had not been tested with DNA sequence data; intraspecific variation in morphological characters diagnostic for the species had not been assessed with specimens from non-type localities, and the species status of the Japanese population(s) was uncertain. We examined the morphology and mitochondrial COI (cytochrome c oxidase subunit I) gene sequences of Capitella specimens from two localities (Ainan and Gamo) in Japan. Specimens from Ainan and Gamo differed from C. teleta from Massachusetts in methyl-green staining pattern, shape of the genital spines, and shape of the capillary chaetae; we concluded that these characters vary intraspecifically. Species delimitation analyses of COI sequences suggested that worms from Ainan and Massachusetts represent C. teleta; these populations share a COI haplotype. The specimens from Gamo may represent a distinct species and comprise a sister group to C. teleta s. str.; we refer to the Gamo population as Capitella aff. teleta. The average Kimura 2-parameter (K2P) distance between C. teleta s. str. and C. aff. teleta was 3.7%. The COI data indicate that C. teleta actually occurs in both the NW Atlantic and NW Pacific. Given the short planktonic larval duration of C. teleta, this broad distribution may have resulted from anthropogenic dispersal.

Detailed reconstruction of the nervous and muscular system of Lobatocerebridae with an evaluation of its annelid affinity.

BACKGROUND: The microscopic worm group Lobatocerebridae has been regarded a 'problematicum', with the systematic relationship being highly debated until a recent phylogenomic study placed them within annelids (Curr Biol 25: 2000-2006, 2015). To date, a morphological comparison with other spiralian taxa lacks detailed information on the nervous and muscular system, which is here presented for Lobatocerebrum riegeri n. sp. based on immunohistochemistry and confocal laser scanning microscopy, supported by TEM and live observations. RESULTS: The musculature is organized as a grid of longitudinal muscles and transverse muscular ring complexes in the trunk. The rostrum is supplied by longitudinal muscles and only a few transverse muscles. The intraepidermal central nervous system consists of a big, multi-lobed brain, nine major nerve bundles extending anteriorly into the rostrum and two lateral and one median cord extending posteriorly to the anus, connected by five commissures. The glandular epidermis has at least three types of mucus secreting glands and one type of adhesive unicellular glands. CONCLUSIONS: No exclusive "annelid characters" could be found in the neuromuscular system of Lobatocerebridae, except for perhaps the mid-ventral nerve. However, none of the observed structures disputes its position within this group. The neuromuscular and glandular system of L. riegeri n. sp. shows similarities to those of meiofaunal annelids such as Dinophilidae and Protodrilidae, yet likewise to Gnathostomulida and catenulid Platyhelminthes, all living in the restrictive interstitial environment among sand grains. It therefore suggests an extreme evolutionary plasticity of annelid nervous and muscular architecture, previously regarded as highly conservative organ systems throughout metazoan evolution.

A new species of Mesochaetopterus (Annelida, Chaetopteridae) from Hong Kong, with comments on the phylogeny of the family.

We described a new species, Mesochaetopterus tingkokensis, based on 14 specimens collected from an intertidal area in Hong Kong. This species is large (body length of complete specimens 17.9-24.1 cm), with 9, 2 and 36-41 chaetigers in region A, B and C, respectively. It belongs to a small group of Mesochaetopterus species with an expanded wing-shaped notopodia in chaetiger B2. The new species can be distinguished from other Mesochaetopterus species in this group by having a pair of palps with two longitudinal stripes formed by suture-like discontinuous orange bands, more teeth in the uncini of region B and C neuropodia, and presence of a bundle of simple chaetae in region C notopodia. Comparison with other chaetopterids based on partial cytochrome oxidase I (COI), 18S and 28S rRNA gene sequences confirmed the placement of M. tingkokensis n. sp.within Mesochaetopterus and its distinction from other members of this genus with the available DNA sequences. The phylogenetic tree base on COI showed that Mesochaetopterus and Chaetopterus are paraphyletic, but that based on concatenated data, 18S and 28S showed they are monophyletic with low supporting values.

Fossilized spermatozoa preserved in a 50-Myr-old annelid cocoon from Antarctica.

The origin and evolution of clitellate annelids--earthworms, leeches and their relatives--is poorly understood, partly because body fossils of these delicate organisms are exceedingly rare. The distinctive egg cases (cocoons) of Clitellata, however, are relatively common in the fossil record, although their potential for phylogenetic studies has remained largely unexplored. Here, we report the remarkable discovery of fossilized spermatozoa preserved within the secreted wall layers of a 50-Myr-old clitellate cocoon from Antarctica, representing the oldest fossil animal sperm yet known. Sperm characters are highly informative for the classification of extant Annelida. The Antarctic fossil spermatozoa have several features that point to affinities with the peculiar, leech-like 'crayfish worms' (Branchiobdellida). We anticipate that systematic surveys of cocoon fossils coupled with advances in non-destructive analytical methods may open a new window into the evolution of minute, soft-bodied life forms that are otherwise only rarely observed in the fossil record.

Ultrastructural study of spermatogenesis and sperm in the African medicinal leech Hirudo troctina Johnson, 1816 (Annelida, Hirudinida).

This paper presents the process of spermatogenesis in the leech Hirudo troctina Johnson, 1816 using light, fluorescent and transmission electron microscopy. At the onset of spermatogenesis in testes, the pear-shaped spermatogonia divide mitotically without full cytokinesis and as a result isogenic groups are formed (clusters, clones) with 2, 4, 8, 16, 32, 64, 128 spermatogonia and, finally, 256 primary spermatocytes occur. The final meiotic divisions of spermatocytes give rise to clones with 1024 spermatids. There are hundreds of developing germ-line clones in each testis. In each clone, the male germ cells divide in full synchrony and they are in the same phase of spermatogenesis. During complex spermiogenesis each spermatid becomes a filiform spermatozoon with a helicoid nucleus, which is characterized by the presence of a long acrosome with two regions - anterior and posterior, which are followed by a helicoid nucleus, a midpiece with only one mitochondrion and a long flagellum. Our results were compared to those on other clitellate annelids that have been studied to date, especially to sperm formation in Hirudo medicinalis Linnaeus, 1785. Only minor differences were found in the length and the diameter of different organelles and the number of spermatids in germ-line clones.

Development and larval feeding in the capitellid annelid Notomastus cf. tenuis.

Making inferences about the evolution of larval nutritional mode and feeding mechanisms in annelids requires data on the form and function of the larvae, but such data are lacking for many taxa. Though some capitellid annelids are known or suspected to have planktotrophic larvae, these larvae have not previously been described in sufficient detail to understand how they feed. Here we describe embryos and larvae of the capitellid Notomastus cf. tenuis from San Juan Island, Washington State. Fertilized oocytes average about 58 µm in equivalent spherical diameter. Early embryos undergo spiral cleavage and develop into larvae that feed for about 5 weeks before metamorphosis. Larvae of N. cf. tenuis capture food particles between prototrochal and metatrochal ciliary bands and transport them to the mouth in an intermediate food groove; this arrangement is typical of "opposed band" larval feeding systems. Surprisingly, however, larvae of N. cf. tenuis appeared to have only simple cilia in the prototrochal ciliary band; among planktotrophic larvae of annelids, simple cilia in the prototroch were previously known only from members of Oweniidae. The anteriormost tier of prototrochal cilia in N. cf. tenuis appears to be non-motile; its role in swimming or particle capture is unclear. Like some planktotrophic larvae in the closely related Echiuridae and Opheliidae, larvae of N. cf. tenuis can capture relatively large particles (up to at least 45 µm in diameter), suggesting that they may use an alternative particle capture mechanism in addition to opposed bands of cilia.

An ultrastructural study of the ovary cord organization and oogenesis in Erpobdella johanssoni (Annelida, Clitellata: Hirudinida).

The aims of the present study were to analyze the ovary cord structure and oogenesis in Erpobdella johanssoni under light, fluorescent and transmission electron microscopy and to compare the obtained results with other clitellate annelids, especially with other arhynchobdellid leeches. Each of the paired ovaries is composed of the ovary wall (ovisac) and several (7-8) short, cone-shaped ovary cords. The ovary cords are of the "Erpobdella" type, i.e. they are short and polarized and five zones containing germ cells at consecutives stages of their development can be distinguished along their long axis. One, huge somatic cell (the apical cell), oogonia and premeiotic germ cells occur at the tip of the apical part of the ovary cord - zone I. Below, in zone II germ cells enter meiosis, whereas in zone III only a few cells continue meiosis and gather nutrients (oocytes), while the rest become nurse cells. In zone IV, huge vitellogenic oocytes form protuberances on the surface of the cord, and degenerating germ cells were observed at the base of the ovary cord (zones IV and V). The germline cells form syncytial cysts in zones I-III. The germline cysts have broadly the same architecture as in the ovaries of all of the clitellate annelids that have been described to date. Each germ cell in a cyst has only one cytoplasmic bridge connecting it to the common cytoplasmic mass - the cytophore. The cytophore is poorly developed, and it has the form of thin, long cytoplasmic strands. The presence of two categories of germ cells suggests a meroistic mode of oogenesis. The germline cysts are closely associated with somatic, follicular cells. There are two subpopulations of follicular cells: one envelops the growing oocytes, while the second is distributed between other germ cells. The entire ovary cord is additionally enveloped by a layer of somatic cells with a spongy appearance - the spongiosa cells. A characteristic feature of vitellogenic oocytes is the condensation of the chromosomes into a karyosome. Fully grown oocytes are excluded from the ovary cords and float freely in the ovisac lumen.

Development and embryonic pattern of body wall musculature in the crassiclitellate Eisenia andrei (Annelida, Clitellata).

During early development of Eisenia andrei (Crassiclitellata), a loose arrangement of primary circular and longitudinal muscles encloses the whole embryo. Circular muscles differentiate in an anterior-posterior progression creating a segmental pattern. Primary circular muscles emerge at the segmental borders while later in development the central part of each segment is filled with circular strands. Longitudinal muscles develop in an anterio-posterior manner as well, but by continuous lengthening. Muscle growth is not restricted by segmental boundaries. The development begins with one pair of prominent longitudinal muscles differentiating ventrally along the right and the left germ band. These first muscles provide a guiding structure for the parallel organization of the afterwards differentiating longitudinal musculature. Additional primary longitudinal muscles emerge and form, together with the initial circular muscles, the primary muscle grid of the embryo. During the following development, secondary longitudinal muscle strands develop and integrate themselves into the primary grid. Meanwhile the primary circular muscles split into thin strands in a ventral to dorsal progression. Thus, a fine structured mesh of circular and longitudinal muscles is generated. Compared to other "Oligochaeta", embryonic muscle patterns in E. andrei are adapted to the development of a lecithotrophic embryo. Nevertheless, two general characteristics of annelid muscle development become evident. The first is the segmental development of the circular muscles from a set of initial muscles situated at the segment borders. Second, there is a continuous development of primary longitudinal muscles starting at the anterior pole. At least one pair of main primary longitudinal strands is characteristic in Annelida. The space between all primary strands is filled with secondary longitudinal strands during further development.

Is Diurodrilus an annelid?

Interstitial marine meiofaunal worms of the genus Diurodrilus have always been considered part of Annelida, either as basal or derived, though generally with reference to Dinophilidae. New evidence shows that Diurodrilus has a unique anatomy, and lacks key annelid features, possibly even segmentation. We assessed the systematic position of Diurodrilus among other protostome animals via light microscopy, confocal laser scanning microscopy, and transmission electron microscopy studies of anatomy, focusing on musculature, the nervous system, as well as molecular sequence data. We show that there is little morphological or molecular evidence to support a relationship with Dinophilidae or any other annelids. Diurodrilus has some similarities to Micrognathozoa, though the latter shows complex jaws. On the basis of the configuration of the nervous system and the cuticle we regard Diurodrilus to belong to Spiralia, possibly close to Annelida; however, until further evidence is acquired it should be regarded as incertae sedis in this large animal clade.

[Ultrastructural characteristics of interneuronal and neuromuscular relations in the trunkal part of the intraepidermal nervous system in Nerilla sp. (Archiannelida)].

This study deals with ultrastructural analysis of interneuronal and neuromuscular relations in a representative of archiannelid Nerilla sp. with primitive intraepidermal type of the nervous system. A particular attention has been paid to the area of ventral ciliate groove and the associated site of epidermis. In the ciliate groove, sensitive and motor cilia are revealed and described. Sites of axonal terminals of the sensitive cells supplied with cilia are noted in the epidermal nervous plexus. Epidermal-muscular cells and nervous terminals on them are revealed. Various interneuronal contact variants both of non-synaptic and of typically synaptic types are described. An attention is drawn to a rare presence of compounds from the type of tight junctions among interneuronal contacts in Nerilla sp. In sufficiently differentiated synapses of the chemical type, phenomena of exocytosis are described. There are shown specific features of innervation of longitudinal (somatic) musculature of the nerillid body with entrance of synaptic vesicles into the basal plate substance and their translocation into the depth of the muscular layer.