Secondary-tail formation during stolonization in the Japanese green syllid, Megasyllis nipponica.

Benthic annelids belonging to the family Syllidae show a distinctive sexual reproduction mode called "stolonization," in which posterior segments are transformed into a reproductive individual-like unit called a "stolon." Megasyllis nipponica forms a stolon head and a secondary tail in the middle of the trunk before a stolon detaches, while, in the case of posterior amputation, posterior regeneration initiates at the wound after amputation. To understand the difference between posterior regeneration and secondary-tail formation during stolonization, detailed comparisons between the developmental processes of these two tail-formation types were performed in this study. Morphological and inner structural observations (i.e., cell proliferation and muscular/nervous development) showed that some processes of posterior regeneration, such as blastema formation and muscular/nervous regeneration at the amputation site, are missing during secondary-tail formation. In contrast, the secondary tail showed some unique features, such as the formation of ventrolateral half-tail buds that later fused in the middle and muscle/nerve branches formed before the detachment of the stolon. These novel features in the process of stolonization are suggested to be adaptive since the animals need to recover a posterior end quickly to stolonize again.

Nerves and availability of mesodermal cells are essential for the function of the segment addition zone (SAZ) during segment regeneration in polychaete annelids.

Most of annelids grow all over their asexual life through the continuous addition of segments from a special zone called "segment addition zone" (SAZ) adjacent to the posterior extremity called pygidium. Amputation of posterior segments leads to regeneration (posterior regeneration-PR) of the pygidium and a new SAZ, as well as new segments issued from this new SAZ. Amputation of anterior segments leads some species to regeneration (anterior regeneration-AR) of the prostomium and a SAZ which produces new segments postero-anteriorly as during PR. During the 1960s and 1970s decades, experimental methods on different species (Syllidae, Nereidae, Aricidae) showed that the function of SAZ depends on the presence and number of mesodermal regeneration cells. Selective destruction of mesodermal regeneration cells in AR had no effect on the regeneration of the prostomium, but as for PR, it inhibited segment regeneration. Thus, worms deprived of mesodermal regeneration cells are always able to regenerate the pygidium or the prostomium, but they are unable to regenerate segments, a result which indicates that the SAZ functions only if these regeneration cells are present during PR or AR. Additionally, during AR, nerve fibres regenerate from the cut nerve cord toward the newformed brain, a situation which deprives the SAZ of local regenerating nerve fibres and their secreted growth factors. In contrast, during PR, nerve fibres regenerate both during the entire regeneration phase and then in normal growth. This review summarizes the experimental evidence for mesoderm cell involvement in segment regeneration, and the differential impact of the digestive tube and the regenerated nerve cord during PR vs AR.

Within-family plasticity of nervous system architecture in Syllidae (Annelida, Errantia).

BACKGROUND: The ground pattern underlying the nervous system of the last common ancestor in annelids was long thought to be settled, consisting of a dorsal brain, circumoesophageal connectives and a subepithelial, ladder-like ventral nerve cord with segmental ganglia connected by paired connectives. With the advent of immunocytochemical stainings and confocal laser scanning microscopy, it becomes evident that its architecture is extremely diverse, which makes the reconstruction of a ground pattern in annelida challenging. Whereas the nervous systems of many different families has already been described, only very few studies looked at the diversity of nervous systems within such clades to give a closer estimate on how plastic the annelid nervous system really is. So far, little is known on syllid nervous system architecture, one of the largest and most diverse groups of marine annelids. RESULTS: The position of the brain, the circumoesophageal connectives, the stomatogastric nervous system, the longitudinal nerves that traverse each segment and the innervation of appendages are relatively uniform within the clade. Both the number of connectives within the ventral nerve cord and the number of segmental nerves, which in earlier studies were used to infer phylogenetic relationships and to reconstruct an annelid ground pattern, are highly diverse and differ between genera or even within a given genus. Differences in the distribution of somata of the brain, the nuchal innervation and its associated cell bodies were found between Syllinae and Exogoninae and may be subfamily-specific. CONCLUSIONS: The nervous system morphology of syllids very likely depends on the taxon-specific ecological requirements. Thus, it is not surprising that in a clade, which occupies such diverse niches as the Annelida, we find similar patterns in phylogenetically widely separated species in similar niches and a high degree of modularity within a family. Only standardized protocols and staining methods can lead to comparable results, but so far different approaches have been taken to describe annelid nervous systems, making homologization of certain structures difficult. This study provides the first thorough description of the nervous system in the family Syllidae, allowing more detailed comparisons between annelid families in the future.

Comparative transcriptomics in Syllidae (Annelida) indicates that posterior regeneration and regular growth are comparable, while anterior regeneration is a distinct process.

BACKGROUND: Annelids exhibit remarkable postembryonic developmental abilities. Most annelids grow during their whole life by adding segments through the action of a segment addition zone (SAZ) located in front of the pygidium. In addition, they show an outstanding ability to regenerate their bodies. Experimental evidence and field observations show that many annelids are able to regenerate their posterior bodies, while anterior regeneration is often limited or absent. Syllidae, for instance, usually show high abilities of posterior regeneration, although anterior regeneration varies across species. Some syllids are able to partially restore the anterior end, while others regenerate all lost anterior body after bisection. Here, we used comparative transcriptomics to detect changes in the gene expression profiles during anterior regeneration, posterior regeneration and regular growth of two syllid species: Sphaerosyllis hystrix and Syllis gracilis; which exhibit limited and complete anterior regeneration, respectively. RESULTS: We detected a high number of genes with differential expression: 4771 genes in S. hystrix (limited anterior regeneration) and 1997 genes in S. gracilis (complete anterior regeneration). For both species, the comparative transcriptomic analysis showed that gene expression during posterior regeneration and regular growth was very similar, whereas anterior regeneration was characterized by up-regulation of several genes. Among the up-regulated genes, we identified putative homologs of regeneration-related genes associated to cellular proliferation, nervous system development, establishment of body axis, and stem-cellness; such as rup and JNK (in S. hystrix); and glutamine synthetase, elav, slit, Hox genes, ß-catenin and PL10 (in S. gracilis). CONCLUSIONS: Posterior regeneration and regular growth show no significant differences in gene expression in the herein investigated syllids. However, anterior regeneration is associated with a clear change in terms of gene expression in both species. Our comparative transcriptomic analysis was able to detect differential expression of some regeneration-related genes, suggesting that syllids share some features of the regenerative mechanisms already known for other annelids and invertebrates.

The Syllis gracilis species complex: A molecular approach to a difficult taxonomic problem (Annelida, Syllidae).

Syllis gracilis is an emblematic member of the subfamily Syllinae (Syllidae, Annelida), which inhabits shallow, temperate coastal waters and can be found on algae, coral rubble, and sponges. Their distinctive ypsiloid chaetae, usually found in specimens from populations all around the world, led to the consideration of the species as cosmopolitan, even though four other species have similar chaetae: Syllis magellanica, S. picta, S. mayeri and S. ypsiloides. The discovery of deeply divergent lineages in the Mediterranean Sea, that were morphologically similar, questioned the cosmopolitanism of S. gracilis and suggested the possibility of it being a species complex. In order to assess the speciation patterns within the putative S. gracilis complex, we undertook species delimitation and phylogenetic analyses on 61 specimens morphologically ascribed to Syllis gracilis and closely related species using a multilocus molecular dataset (two mitochondrial and two nuclear markers). Our results suggest high levels of genetic differentiation between the S. gracilis populations analyzed, some of which have morphologically distinctive features. Five to eight distinct lineages (depending on the analysis) were identified, all with geographically restricted distributions. Although the presence of ypsiloid chaetae has been traditionally considered the main character to identify S. gracilis, we conclude that this feature is homoplastic. Instead, we propose that characters such as the degree of fusion of blades and shafts in chaetae, the morphology of the posterior chaetae or the animal color pattern should be considered to differentiate lineages within the S. gracilis species complex. Our study does not support the cosmopolitanism of S. gracilis, and instead provides morphological and molecular evidence of the existence of a complex of pseudo-cryptic species.

A new genus and species of nudibranch-mimicking Syllidae (Annelida, Polychaeta).

Nudibranch mollusks, which are well-known for their vivid warning coloration and effective defenses, are mimicked by diverse invertebrates to deter predation through both Müllerian and Batesian strategies. Despite extensive documentation across different taxa, mimickers have not been detected among annelids, including polychaetes, until now. This study described a new genus and species of polychaete living on Dendronephthya octocorals in Vietnam and Japan. Belonging to Syllidae, it exhibits unique morphological adaptations such as a low number of body segments, simple chaetae concealed within the parapodia and large and fusiform antennae and cirri. Moreover, these appendages are vividly colored, featuring an internal dark red area with numerous terminal white spots and bright yellow tips, effectively contributing to mimicking the appearance of a nudibranch. This discovery not only documents the first known instance of such mimicry among annelids, but also expands our understanding of evolutionary adaptation and ecological strategies in marine invertebrates.

Three new species of Syllis Savigny in Lamarck, 1818 (Annelida, Syllidae) from Chilean Patagonia.

Three new species of the genus Syllis Savigny in Lamarck, 1818, from the Chilean Patagonia are described, figured and discussed. The specimens were collected in boulders, sediment bottoms, Macrocystis pyrifera holdfasts, and inside tubes of Chaetopterus cf. variopedatus. Syllis patagonica n. sp., is characterized by its color pattern, short, fusiform dorsal cirri, plenty of hyaline inclusions, acuminate posterior acicula, and compound chaetae with slightly bidentate falcigers. Syllis terraeignium n. sp., has similar compound chaetae, but dorsal cirri are longer than those of the above mentioned species, a long proventricle, and the posterior acicula are straight, pointed. Finally, Syllis patersoni n. sp., has a slender, elongated body, with moderately long dorsal cirri, plenty of hyaline inclusions, acuminate posterior acicula, and compound chaetae composed by falcigers and short spiniger-like.

Sex-specific gene expression differences in reproducing Syllis prolifera and Nudisyllis pulligera (Annelida, Syllidae).

Syllidae is an annelid family characterized by its complex life cycles involving some of the most outstanding annelid reproductive strategies. Syllid reproductive modes sometimes imply the modification of the posterior body to form independent reproductive units (schizogamy) or the development of swimming adults (epigamy). These modes of sexual reproduction have been studied for more than 150 years, and yet, little is known regarding their molecular background. Notably, while several studies during the last three decades have revealed details about molecular mechanisms involved in the reproduction of some few model annelids, studies focusing on syllids remain limited. Thus, we performed differential gene expression analyses of female, male, and non-reproducing individuals of Syllis prolifera (schizogamic) and Nudisyllis pulligera (epigamic), as representatives of two different reproductive strategies. For that, transcriptomes from specimens of three conditions (non-reproducing, male, female) were de novo assembled and annotated for S. prolifera and N. pulligera. We found rather similar gene expression profiles for female and non-reproducing individuals, while male gene expression is clearly different. Although previous studies have suggested that femininity in syllids might require additional signalling, our analyses support a scenario, where masculinity may also involve several specific genetic processes.

An integrative approach to the anatomy of Syllis gracilis Grube, 1840 (Annelida) using micro-computed X-ray tomography.

BACKGROUND: The overall anatomy of the genus Syllis (Annelida: Syllidae) has been largely studied; however, an integrative approach considering different anatomical techniques has never been considered. Here, we use micro-computed X-ray tomography (micro-CT) to examine the internal anatomy of Syllis gracilis Grube, 1840, along with other widely available techniques. METHODS: We studied the anatomy of the marine annelid S. gracilis through an integrative approach, including micro-CT along with stereo and light compound microscopy (STM, LCM), scanning electron microscopy (SEM), confocal laser scanning microscopy (CLSM) and histological sectioning (HIS). In this manner, we evaluated the applicability of micro-CT for the examination of annelid anatomy by testing whether the images obtained make it possible to visualize the main body structures, in comparison with other current techniques, of the various elements of its internal anatomy. RESULTS: Overall external and internal body elements are clearly shown by the integrative use of all techniques, thus overcoming the limitations of each when studied separately.Any given method shows disparate results, depending on the body part considered. For instance, micro-CT provided good images of the external anatomy, including relevant characters such as the shape, length and number of articles of dorsal parapodial cirri. However, it is especially useful for the examination of internal anatomy, thus allowing for 3D visualization of the natural spatial arrangement of the different organs. The features best visualized are those of higher tissue density (i.e., body musculature, anterior parts of the digestive tract), particularly in 3D images of unstained specimens, whereas less electrodense tissues (i.e., the peritoneal lining of septa and nervous system) are less clearly visualized. The use of iodine stain with micro-CT has shown advantages against non-staining for the adequate observation of delicate elements of low density, such as the segmental organs, the connective between the ganglia, the ventral nerve cord and segmental nerves. DISCUSSION: Main external anatomical elements of S. gracilis are well shown with micro-CT, but images show lesser optical resolution and contrast when compared to micrographs provided by SEM and CLSM, especially for fine structural features of chaetae. Comparison of micro-CT and HIS images revealed the utility and reliability of the former to show the presence, shape and spatial disposition of most internal body organs; the resolution of micro-CT images at a cellular level is, however, much lower than that of HIS, which makes both techniques complementary.

Syllidae (Annelida: Phyllodocida) from the deep Mediterranean Sea, with the description of three new species.

Despite almost two centuries of research, the diversity of Mediterranean deep-sea environments remain still largely unexplored. This is particularly true for the polychaete family Syllidae. We report herein 14 species; among them, we describe Erinaceusyllis barbarae n. sp., Exogone sophiae n. sp. and Prosphaerosyllis danovaroi n. sp. and report Parexogone wolfi San Martín, 1991, Exogone lopezi San Martín, Ceberio Aguirrezabalaga, 1996 and Anguillosyllis Day, 1963 for the first time from the Western Mediterranean, the latter based on a single individual likely belonging to an undescribed species. Moreover, we re-establish Syllis profunda Cognetti, 1955 based on type and new material. Present data, along with a critical analysis of available literature, show that Syllidae are highly diverse in deep Mediterranean environments, even though they are rarely reported, probably due to the scarce number of studies devoted to the size-fraction of benthos including deep-sea syllids. Most deep-sea Syllidae have wide distributions, which do not include shallow-waters. 100 m depth apparently represents the boundary between the assemblages dominated by generalist shallow water syllids like Exogone naidina Ørsted, 1843 and Syllis parapari San Martín López, 2000, and those deep-water assemblages characterised by strictly deep-water species like Parexogone campoyi San Martín, Ceberio Aguirrezabalaga, 1996, Parexogone wolfi San Martín, 1991 and Syllis sp. 1 (= Langerhansia caeca Katzmann, 1973).

The genus Syllis Savigny in Lamarck, 1818 (Annelida: Syllidae: Syllinae) from Australia (second part): four new species and re-description of twelve previously described species.

Syllis Savigny in Lamarck, 1818, the type genus of Syllidae, is the largest and most diverse within the family. This genus presents many taxonomic and phylogenetic problems due mainly to the lack of molecular data and morphological synapomorphies for many of the species, but also to poor or inadequate descriptions. In order to improve the knowledge of the genus, we have undertaken a morphological revision of Australian species, based on type material and material from the Australian Museum. Sixteen species are herein described, of which four are new and twelve are redescriptions of alreadyknown species, with six new combinations: Syllis boggemanni, n. sp.; S. joaoi, n. sp.; S. karlae, n. sp.; S. marceloi, n. sp.; S. albanyensis (Hartmann-Schröder, 1984) n. comb.; S. erikae (Hartmann-Schröder, 1981) n. comb.; S. krohnii Ehlers 1864; S. lunaris (Imajima, 1966) n. comb.; S. lutea (Hartmann-Schröder, 1960); S. macrodentata (Hartmann-Schröder, 1982) n. comb.; S. monilaris Savigny in Lamarck, 1818; S. nigropunctata Haswell, 1886; S. pectinans Haswell, 1920; S. rosea (Langerhans, 1879); S. warrnamboolensis (Hartmann-Schröder, 1987) n. comb.; and S. yallingupensis (Hartmann-Schröder, 1982), n. comb.

Exogoninae (Annelida: Syllidae) from Chilean Patagonia.

The subfamily Exogoninae was studied from samples collected in shallow waters of the fjords and channels of the Patagonian region of Chile. Two new species are described: Exogone yagan n. sp. and Erinaceusyllis carrascoi n. sp. The species Exogone heterosetoides, Erinaceusyllis bidentata and Erinaceusyllis perspicax are newly reported to Chile, as well as the genus Erinaceusyllis San Martín, 2005. Parapionosyllis brevicirra, Sphaerosyllis hirsuta and Salvatoria rhopalophora, n. comb., are also reported, with the latter redescribed. Finally, we redescribe Exogone anomalochaeta from Antarctica. Most of the species were found inside tubes of Chaetopterus cf. variopedatus; this habitat is new for Exogoninae. This research is a new taxonomic account of Syllidae in Chile and improves the knowledge of Exogoninae of the Patagonian region.

Taxonomic contribution to the genus Sphaerosyllis (Annelida: Syllidae: Exogoninae) in the Black Sea.

During a study of fauna associated with a shallow-water Zostera (Zosterella) noltei bed from the southern part of the Romanian Black Sea coast, among the identified material collected in 2015, a syllid polychaete belonging to the subfamily Exogoninae, Sphaerosyllis taylori Perkins, 1981, represents a new record for the Black Sea. Re-examination of available specimens previously identified as Sphaerosyllis bulbosa Southern, 1914 revealed that they belong to an unknown species, described herein as Sphaerosyllis pontica sp. nov. The new species is characterized by the median antenna inserted more posteriorly than the lateral antennae, dorsal cirri with bulbous bases and very short tips, shorter than the parapodial lobes, dorsal cirri absent on chaetiger 2, parapodial glands with fibrillar material from chaetiger 4 onwards, compound chaetae with short blades and smooth shafts, anterior parapodia with two aciculae each, one straight and one with bent tip. Descriptions of both species are provided together with a key to all Sphaerosyllis species known from the Black Sea.

Syllidae (Annelida, Polychaeta) from the Caribbean coast of Venezuela.

Venezuela possesses a great variety of coastal environments allowing for a high diversity of marine species. However, systematic studies on marine invertebrates are scarce, especially on polychaetes. The family Syllidae is poorly known, and only 14 genera and 42 species have been reported from this country. A total of 13 genera and 26 species the Syllidae were identified from benthic samples collected on different substrata of the northeastern coast of Venezuela. Of these, seven genera and 16 species constitute new records for Venezuela: Odontosyllis guillermoi, Syllides floridanus, Salvatoria clavata, Salvatoria limbata, Sphaerosyllis longicauda, Parapionosyllis longicirrata, Trypanosyllis parvidentata, Trypanosyllis vittigera, Opisthosyllis sp., Syllis amica, Syllis armillaris, Syllis gracilis, Syllis pseudoarmillaris, Syllis vittata, Parasphaerosyllis indica and Myrianida convoluta.

Sphaerosyllis levantina sp. n. (Annelida) from the eastern Mediterranean, with notes on character variation in Sphaerosyllis hystrix Claparède, 1863.

Examination of polychaete specimens from Haifa Bay (Israel, eastern Mediterranean Sea) revealed several individuals exhibiting morphological characteristics similar to Sphaerosyllis hystrix Claparède, 1863. A detailed morphometrical analysis of the Israeli specimens in comparison to specimens of Sphaerosyllis hystrix and Sphaerosyllis boeroi Musco, Çinar and Giangrande, 2005 supported the description of the former as a new species, Sphaerosyllis levantinasp. n. Individuals of Sphaerosyllis hystrix formed a very heterogeneous group with strong character variations in the analysis and the presumed cosmopolitan distribution of the species is discussed based on literature records.

An account of the taxonomy and distribution of Syllidae (Annelida, Polychaetes) in the eastern Mediterranean, with notes on the genus Prosphaerosyllis San Martín, 1984 in the Mediterranean.

The syllid fauna of three locations in Crete and Israel (eastern Mediterranean Sea) was studied, yielding 82 syllid species, many of which were found for the first time in the respective areas: Seventeen species were recorded for the first time on the Israeli coasts and 20 in Greek waters. Perkinsyllis augeneri (Hartmann-Schröder, 1979) and Prosphaerosyllis chauseyensis Olivier et al., 2011 are new records for the Mediterranean Sea. Detailed information is given on the morphology, ecology and distribution of the species recorded for the first time in the studied areas. In addition, an update on the distribution of the genus Prosphaerosyllis San Martín, 1984 in the Mediterranean is given and an identification key to the Mediterranean species is provided.

Sílidos (Syllidae: Polychaeta) del Parque Nacional de Coiba (Pacífico, Panamá) / Syllids (Syllidae: Polychaeta) from Coiba National Park (Pacific of Panama)

During a study carried out on soft bottoms from Coiba National Park (Panama), 218 specimens of syllids (Annelida: Polychaeta) belonging to 19 interstitial species have been identified. Two species are new reports for the Pacific Ocean, Exogone (Exogone) arenosa Perkins, 1981 and Streptosyllis websteri Southern, 1914; five are new for Panama, E. (E.) dispar (Webster, 1879); E. (E.) longicornis Westheide, 1974; Salvatoria mediodentata (Westheide, 1974); Pionosyllis heterocirrata (Hartmann-Schrõder, 1959) and Syllis glarearia (Westheide, 1974). A characteristic not mentioned in the original description is herein reported for E. (E.) longicornis: the presence of triangular subterminal processes in the spiniger-like compound chaetae of chaetiger one. Specimens of the species Syllis botosaneanui Hartmann-Schrõder, 1973 harboring inside embryos of various stages of development have been found; this is the first report for this species as viviparous. Lastly, one specimen of the genus Syllis has been found that is mainly characterized by its long pharynx, two dorsal prostomial lobes, and compound chaetae with short blades and long spinulation. Because we consider that one specimen is not sufficient to describe a new species we refer it as Syllis sp.

Durante un estudio sobre la fauna marina del Parque Nacional de Coiba (Panamá) se identificaron 281 ejemplares de sílidos (Annelida: Polychaeta) intersticiales pertenecientes a 19 especies. Dos son nuevas citas para el Pacífico, Exogone (Exogone) arenosa Perkins, 1981 y Streptosyllis websteri Southern, 1914; cinco son nuevas citas para Panamá, E. (E.) dispar (Webster, 1879), E. (E.) longicornis Westheide, 1974, Salvatoria mediodentata (Westheide, 1974); Pionosyllis heterocirrata (Hartmann-Schröder, 1959) y Syllis glarearia (Westheide, 1974). Se describe un caracter no presente en la descripción original de E. Longicornis, la presencia de procesos triangulares subterminales en la primera pseudospinígera. Se describen ejemplares de la especie Syllis botosaneanui Hartmann-Schröder, 1973 con embriones en diverso estado de desarrollo en su interior, por lo que se trata de la primera referencia de esta especie como vivípara. Por último, se describe un ejemplar de Syllis sp., que se caracteriza fundamentalmente por la posesión de una larga faringe, dos lóbulos prostomiales dorsales y por sus sedas compuestas de artejos cortos y larga espinulación. Estas características diferencian a Syllis sp. de cualquier especie del género pero no se describe como especie nueva por disponerse únicamente de un solo ejemplar.