New fossil of Gaoloufangchaeta advances the origin of Errantia (Annelida) to the early Cambrian.

Molecular clock estimates suggest the origin of Annelida dates back to the Ediacaran period, which is in discordance with the first appearance of this taxon in the early Cambrian, as evidenced by the fossil records of stem-group and basally branching crown-group annelids. Using new material from the early Cambrian Guanshan biota (Cambrian Series 2, Stage 4), we re-interpret Gaoloufangchaeta bifurcus Zhao, Li & Selden, 2023, as the earliest known errantian annelid. Gaoloufangchaeta has a prominent anterior end bearing three pairs of putatively sensory appendages and a pair of anterior eyes; a muscular eversible pharynx with papillae is identified. The presence of enlarged parapodia with acicula-like structures and long capillary chaetae suggests a pelagic lifestyle for this taxon. Our phylogenetic analyses recover Gaoloufangchaeta within the Phyllodocida (Pleistoannelida, Errantia), extending the origin of Errantia back to the early Cambrian. Our data are in line with the hypothesis that Annelida diverged before the Cambrian and indicate both morphological and ecological diversification of annelids in the early Cambrian.

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.

Morphological, histological and gene-expression analyses on stolonization in the Japanese Green Syllid, Megasyllis nipponica (Annelida, Syllidae).

Benthic annelids belonging to the family Syllidae (Annelida, Errantia, Phyllodocida) exhibit a unique reproduction mode called "schizogamy" or "stolonization", in which the posterior body part filled with gametes detaches from the original body, as a reproductive unit (stolon) that autonomously swims and spawns. In this study, morphological and histological observations on the developmental processes during stolonization were carried out in Megasyllis nipponica. Results suggest that the stolon formation started with maturation of gonads, followed by the formation of a head ganglion in the anteriormost segment of the developing stolon. Then, the detailed stolon-specific structures such as stolon eyes and notochaetae were formed. Furthermore, expression profiles of genes involved in the anterior-posterior identity (Hox genes), head determination, germ-line, and hormone regulation were compared between anterior and posterior body parts during the stolonization process. The results reveal that, in the posterior body part, genes for gonadal development were up-regulated, followed by hormone-related genes and head-determination genes. Unexpectedly, Hox genes known to identify body parts along the anterior-posterior axis showed no significant temporal expression changes. These findings suggest that during stolonization, gonad development induces the head formation of a stolon, without up-regulation of anterior Hox genes.

New insights into phylogenetic relationships of Rhabdocoela (Platyhelminthes) including members of Mariplanellida.

BACKGROUND: Previous flatworm phylogenetic research has been carried out analysing 18S and 28S DNA markers. Through this methodology, Mariplanellinae subfamily has been recently re-classified as Mariplanellida status novus. This new classification implied that 3 genera belonged to Mariplanellida: Mariplanella, Lonchoplanella and Poseidoplanella. In this study, we aim to clarify some of the relationships within Rhabdocoela analysing 18S and 28S DNA markers of a total of 91 species through Maximum Likelihood and Bayesian Inference methodologies. A total of 11 species and genera, including Lonchoplanella, from the island of Sylt are included and had not previously been involved in any molecular phylogenetic analyses. RESULTS: Our phylogenetic results support Mariplanellida as an independent group within Rhabdocoela and its status as an infraorder. Our study suggests that Lonchoplanella axi belongs to Mariplanellida. Within Rhabdocoela, Haloplanella longatuba is nested within Thalassotyphloplanida, instead of Limnotyphloplanida. Within Kalyptorhynchia, the taxon Eukalyptorhynchia turned out to be paraphyletic including members of Schizorhynchia. These results also support the position of the genus Toia separate from Cicerinidae. CONCLUSIONS: Lonchoplanella axi belongs to Mariplanellida, whose status as infraorder is herein confirmed. The genus Toia belongs separate from Cicerinidae. Further research is needed to clarify the phylogenetic relationships of Hoploplanella. Most of the species, genera and families included in this study with more than one terminal are monophyletic and well supported. Adding gene markers and complementary morphological studies will help to clarify those relationships that remain uncertain.

Another piece for the syllid puzzle: A new species from Japan and its mitochondrial genome reveal the enigmatic Clavisyllis (Phyllodocida: Syllidae) as a member of Eusyllinae.

The phylogenetic relationships of Syllidae have been analyzed in several studies during the last decades, resulting in highly congruent topologies. Most of the subfamilies were found to be monophyletic, while other groups (Eusyllinae and several genera) have been reorganized attending their phylogenetic relationships. However, there are still several enigmatic genera, which could not be assigned to any of the established subgroups. These enigmatic genera usually show a combination of characters indicating relationships with several different groups, and some show morphological traits unique to Syllidae. One of the most intriguing genera, still unclassified within Syllidae is Clavisyllis Knox. Herein, we provide a complete description of a new species Clavisyllis tenjini n. sp. from Japan. We sequence the complete mitochondrial genome, compare with the available data from other syllids, and perform a phylogenetic analysis of three genes (18S, 16S, COI), traditionally used in previous studies. Clavisyllis shows a unique combination of characters within Syllidae, such as nuchal lappets and large ovoid dorsal cirri. The new species has additional anterior appendages that have not been found in any other syllid. Our results show the genus is a member of Eusyllinae, closely related to Pionosyllis Malmgren. The mitochondrial gene order agrees with the considered plesiomorphic gene order in Annelida, which is present in all members of Eusyllinae investigated so far. Clavisyllis reproduces by epigamy, the reproductive mode of members of Eusyllinae. The present study contributes to the systematics of Syllidae, a complex group with a large number of species and striking reproductive modes.

Trophic ecology of three marine polychaete species: Evidence from laboratory experiments using stable isotope (15N, 13C), fatty acid (NLFA) analyses, and C and N stoichiometry.

Marine polychaetes are a highly diverse taxon with a large variety of different feeding modes. A proper identification of the different diets of polychaete species and their trophic position in the food web is crucial for understanding their interactions in marine ecosystems as well as marine community dynamics. Since gut content analyses and feeding experiments are difficult to conduct in marine habitats, other methods using biochemical tracers may contribute to a better understanding of the trophic ecology of marine polychaetes. Here, we conducted laboratory experiments and used a combination of stable isotope (13C, 15N) and neutral lipid fatty acid (NLFA) analyses, as well as C and N stoichiometry to determine the feeding habits of widely distributed marine polychaete species, such as Eurythoe complanata, Platynereis massiliensis, and Syllis malaquini. Additionally, the impact of starvation on the stable isotope signatures was analyzed. Our data show that the trophic ecology of the three species differs from each other. Stable isotope and fatty acid analyses indicate that (a) E. complanata is mainly feeding on algae, cyanobacteria and to a lesser extent on fungi, that (b) S. malaquini is mainly feeding on algae and bacteria and that (c) P. massiliensis is mainly feeding on algae, especially diatoms. An analysis of the C:N ratio of the respective annelid species and their potential food sources corroborated these results. The combination of stable isotope signatures, fatty acid markers and stoichiometry of carbon and nitrogen is a useful tool to identify the diet and trophic position of marine polychaete species and provides more results about their feeding habits and their position in marine food webs.

First mitochondrial genomes of Chrysopetalidae (Annelida) from shallow-water and deep-sea chemosynthetic environments.

Among Annelida, Chrysopetalidae is an ecologically and morphologically diverse group, which includes shallow-water, deep-sea, free-living, and symbiotic species. Here, the four first mitochondrial genomes of this group are presented and described. One of the free-living shallow-water species Chrysopetalum debile (Chrysopetalinae), one of the yet undescribed free-living deep-sea species Boudemos sp., and those of the two deep-sea bivalve endosymbionts Craseoschema thyasiricola and Iheyomytilidicola lauensis (Calamyzinae). An updated phylogeny of Chrysopetalidae is performed, which supports previous phylogenetic hypotheses within Chrysopetalinae and indicates a complex ecological evolution within Calamyzinae. Additionally, analyses of natural selection pressure in the four mitochondrial genomes and additional genes from the two shallow-water species Bhawania goodei and Arichlidon gathofi were performed. Relaxed selection pressure in the mitochondrion of deep-sea and symbiotic species was found, with many sites under selection identified in the COX3 gene of deep-sea species.

Monsters reveal patterns: bifurcated annelids and their implications for the study of development and evolution.

During recent decades, the study of anatomical anomalies has been of great relevance for research on development and its evolution. Yet most animal groups have never been studied under this perspective. In annelids, one of the most common and remarkable anomalies is anteroposterior axis bifurcation, that is animals that have two or more heads and/or tails. Bifurcated annelids were first described in the 18th century and have been occasionally reported since then. However, these animals have rarely been considered other than curiosities, one-off anomalies, or monsters, and a condensed but comprehensive analysis of this phenomenon is lacking. Such an analysis of the existing knowledge is necessary for addressing the different patterns of annelid bifurcation, as well as to understand possible developmental mechanisms behind them and their evolution. In this review we summarize reports of annelid bifurcation published during the last 275 years and the wide variety of anatomies they present. Our survey reveals bifurcation as a widespread phenomenon found all over the annelid tree. Moreover, it also shows that bifurcations can be classified into different types according to anatomy (lateral versus dorsoventral) or developmental origin (embryonic versus postembryonic, the latter occurring in relation to regeneration, reproduction, or growth). Regarding embryos, three different types of bifurcation can be found: conjoined twins (in clitellates); Janus embryos (two posterior ends with a single head which shows duplicated structures); and duplicitas cruciata embryos (with anterior and posterior bifurcation with a 90° rotation). In adults, we show that while lateral bifurcation can result in well-integrated phenotypes, dorsoventral bifurcation cannot since it requires the discontinuity of at least some internal organs. The relevance of this distinction is highlighted in the case of the Ribbon Clade, a group of syllid annelids in which some species reproduce by collateral and successive gemmiparity (which involves dorsoventral bifurcation), while others grow by branching laterally. Although most known cases of bifurcation came from accidental findings in the wild or were unintentionally produced, experimental studies resulting in the induction of bifurcation of both embryos and adults are also reviewed. In embryos, these experimental studies show how mechanical or chemical disruption of the zygote can result in bifurcation. In adults, the ventral nervous system and the digestive tract seem to play a role in the induction of bifurcation. Based on the reviewed evidence, we argue that the long-forgotten study of annelid developmental anomalies should be incorporated into the growing field of annelid EvoDevo and examined with modern techniques and perspectives.

Effects of GSK3ß inhibition in the regeneration of Syllis malaquini (Syllidae, Annelida).

The Wnt/ß-catenin signaling pathway has been widely associated to the reestablishment of anteroposterior body polarities in the embryonic development and regeneration in animals. For instance, in annelids, cellular proliferation, wound healing, and blastema development can be affected when this pathway is disrupted. However, very little is known about the genetic regulatory processes involved in these anomalies. Here, we investigate the morphological effects of 1-azakenpaullone, a pharmacological inhibitor of GSK3ß that is supposed to over-activate the Wnt/ß-catenin pathway, during the anterior and posterior regeneration of the annelid Syllis malaquini. The results showed that high concentrations of 1-azakenpaullone affect the stages of blastema differentiation and resegmentation. Therefore, GSK3ß-associated gene regulatory networks are candidate to investigate the genetic mechanisms involved in the regular course of S. malaquini regeneration.

Cellular proliferation dynamics during regeneration in Syllis malaquini (Syllidae, Annelida).

BACKGROUND: In syllids (Annelida, Syllidae), the regenerative blastema was subject of many studies in the mid and late XXth century. This work on syllid regeneration showed that the blastema is developed by a process of dedifferentiation of cells near the wound, followed by their proliferation and redifferentiation (cells differentiate to the original cell type) or, in some specific cases, transdifferentiation (cells differentiate to a cell type different from the original). Up to date, participation of stem cells or pre-existing proliferative cells in the blastema development has never been observed in syllids. This study provides the first comprehensive description of Syllis malaquini's regenerative capacity, including data on the cellular proliferation dynamics by using an EdU/BrdU labelling approach, in order to trace proliferative cells (S-phase cells) present before and after operation. RESULTS: Syllis malaquini can restore the anterior and posterior body from different cutting levels under experimental conditions, even from midbody fragments. Our results on cellular proliferation showed that S-phase cells present in the body before bisection do not significantly contribute to blastema development. However, in some specimens cut at the level of the proventricle, cells in S-phase located in the digestive tube before bisection participated in regeneration. Also, our results showed that nucleus shape allows to distinguish different types of blastemal cells as forming specific tissues. Additionally, simultaneous and sequential addition of segments seem to occur in anterior regeneration, while only sequential addition was observed in posterior regeneration. Remarkably, in contrast with previous studies in syllids, sexual reproduction was not induced during anterior regeneration of amputees lacking the proventricle, a foregut organ widely known to be involved in the stolonization control. CONCLUSIONS: Our findings led us to consider that although dedifferentiation and redifferentiation might be more common, proliferative cells present before injury can be involved in regenerative processes in syllids, at least in some cases. Also, we provide data for comparative studies on resegmentation as a process that differs between anterior and posterior regeneration; and on the controversial role of the proventricle in the reproduction of different syllid lineages.

Integrative anatomical study of the branched annelid Ramisyllis multicaudata (Annelida, Syllidae).

The sponge-dwelling Syllidae Ramisyllis multicaudata and Syllis ramosa are the only annelid species for which a branched body with one head and multiple posterior ends is known. In these species, the head is located deep within the sponge, and the branches extend through the canal system of their host. The morphology of these creatures has captivated annelid biologists since they were first discovered in the late XIXth century, and their external characteristics have been well documented. However, how their branched bodies fit within their symbiotic host sponges and how branches translate into internal anatomy has not been documented before. These features are crucially relevant for understanding the body of these animals, and therefore, the aim of this study was to investigate these aspects. In order to assess these questions, live observation, as wells as histology, immunohistochemistry, micro-computed tomography, and transmission electron microscopy techniques were used on specimens of R. multicaudata. By using these techniques, we show that the complex body of R. multicaudata specimens extends greatly through the canal system of their host sponges. We demonstrate that iterative external bifurcation of the body is accompanied by the bifurcation of the longitudinal organ systems that are characteristic of annelids. Additionally, we also highlight that the bifurcation process leaves an unmistakable fingerprint in the form of newly-described "muscle bridges." These structures theoretically allow one to distinguish original and derived branches at each bifurcation. Last, we characterize some of the internal anatomical features of the stolons (reproductive units) of R. multicaudata, particularly their nervous system. Here, we provide the first study of the internal anatomy of a branched annelid. This information is not only crucial to deepen our understanding of these animals and their biology, but it will also be key to inform future studies that try to explain how this morphology evolved.

Life Cycle of the Japanese Green Syllid, Megasyllis nipponica (Annelida: Syllidae): Field Collection and Establishment of Rearing System.

Some polychaete species in the family Syllidae exhibit distinctive life cycles, in which a posterior part of the body of an individual detaches as a reproductive individual called a "stolon". This type of reproductive mode is known as stolonization or schizogamy. Although a number of observations have been reported, and techniques using molecular markers have recently been applied to characterize this phenomenon, little is known about the developmental and physiological mechanisms underlying stolonization. In the present study, Megasyllis nipponica, a common syllid species distributed throughout Japan, is proposed as a model to reveal the developmental and physiological mechanism of stolonization, and the rearing system to maintain it in laboratory conditions is described. This species was repeatedly sampled around Hokkaido, where more dense populations were found from August to October. The animals were maintained in the laboratory under stable long-day condition (20°C, 16L:8D), and fed mainly with spinach powder. Stolonization processes, spawning, embryonic and postembryonic development were observed and documented, and the required period of time for each developmental stage was recorded. The complete generation time was around two months under the rearing condition. The information provided is valuable to maintain this and other syllid species in the laboratory, and hence contributes to the establishment of new evolutionary and developmental research lines in this group of annelids.

Expression of vasa, piwi, and nanos during gametogenesis in Typosyllis antoni (Annelida, Syllidae).

Although model species have proven to be crucial for developmental biology, the evo-devo approach requires a broader picture across phylogeny. Herein, we try to expand the range of studied annelids by presenting a transcriptome of Typosyllis antoni as a tool for the study of developmental and evolutionary processes in Syllidae. Moreover, we provide homologs of the stem-cell markers vasa, piwi, and nanos, and investigate their expression patterns in gamete-producing individuals for the first time in this group. We found no expression in females, while there is a distinct expression pattern in males. Based on this data, we argue that spermatogenesis starts in the gonads and finishes in the coelomic cavity, and it occurs simultaneously in a large number of segments. Surprisingly, no expression of the stem-cell markers was found in the segment addition zone of these reproducing animals (stolonizing). Preliminary explanations like a lack of growth during stolonization, or the absence of a common genetic program between germ and somatic stem cells, are discussed. Finally, no reservoir of primordial cells has been detected, suggesting a possible epigenic origin of the Primordial Germ Cells of this species, though this hypothesis needs to be further investigated.

The transcriptome of the Bermuda fireworm Odontosyllis enopla (Annelida: Syllidae): A unique luciferase gene family and putative epitoky-related genes.

The Bermuda fireworm Odontosyllis enopla exhibits an extremely tight circalunar circadian behavior that results in an impressive bioluminescent mating swarm, thought to be due to a conventional luciferase-mediated oxidation of a light-emitting luciferin. In addition, the four eyes become hypertrophied and heavily pigmented, and the nephridial system is modified to store and release gametes and associated secretions. In an effort to elucidate transcripts related to bioluminescence, circadian or circalunar periodicity, as well as epitoky-related changes of the eyes and nephridial system, we examined the transcriptomic profile of three female O. enopla during a bioluminescent swarm in Ferry Reach, Bermuda. Using the well-characterized luciferase gene of the Japanese syllid Odontosyllis undecimdonta as a reference, a complete best-matching luciferase open reading frame (329 amino acids in length) was found in all three individuals analyzed in addition to numerous other paralogous sequences in this new gene family. No photoproteins were detected. We also recovered a predicted homolog of 4-coumarate-CoA ligase (268 amino acids in length) that best matched luciferase of the firefly Luciola with the best predicted template being the crystal structure of luciferase for Photinus pyralis, the common eastern firefly. A wide variety of genes associated with periodicity were recovered including predicted homologs of clock, bmal1, period, and timeless. Several genes corresponding to putative epitoky-related changes of the eyes were recovered including predicted homologs of a phototransduction gene, a retinol dehydrogenase and carotenoid isomerooxygenase as well as a visual perception related retinal rod rhodopsin-sensitive cGMP 3',5'-cyclic phosphodiesterase. Genes correlating to putative epitoky-related changes of the nephridia included predicted homologs of nephrocystin-3 and an egg-release sex peptide receptor.

Regeneration mechanisms in Syllidae (Annelida).

Syllidae is one of the most species-rich groups within Annelida, with a wide variety of reproductive modes and different regenerative processes. Syllids have striking ability to regenerate their body anteriorly and posteriorly, which in many species is redeployed during sexual (schizogamy) and asexual (fission) reproduction. This review summarizes the available data on regeneration in syllids, covering descriptions of regenerative mechanisms in different species as well as regeneration in relation to reproductive modes. Our survey shows that posterior regeneration is widely distributed in syllids, whereas anterior regeneration is limited in most of the species, excepting those reproducing by fission. The latter reproductive mode is well known for a few species belonging to Autolytinae, Eusyllinae, and Syllinae. Patterns of fission areas have been studied in these animals. Deviations of the regular regeneration pattern or aberrant forms such as bifurcated animals or individuals with multiple heads have been reported for several species. Some of these aberrations show a deviation of the bilateral symmetry and antero-posterior axis, which, interestingly, can also be observed in the regular branching body pattern of some species of syllids.

On the role of the proventricle region in reproduction and regeneration in Typosyllis antoni (Annelida: Syllidae).

BACKGROUND: Syllids are a species rich annelid family possessing remarkable regenerative ability, which is not only the response after traumatic injury, but also a key step during the life cycle of several syllid taxa. In these animals the posterior part of the body becomes an epitoke and is later detached as a distinct unit named stolon. Such a sexual reproductive mode is named schizogamy or stolonization. The prostomium and the proventricle, a modified foregut structure, have been proposed to have a control function during this process, though the concrete mechanisms behind it have never been elucidated. RESULTS: By using different experimental set-ups, histology and immunohistochemistry combined with subsequent cLSM analyzes, we investigate and document the regeneration and stolonization in specimens of Typosyllis antoni that were amputated at different levels throughout the antero-posterior body axis. The removal of the anterior end including the proventricle implies an incomplete anterior regeneration as well as severe deviations from the usual reproductive pattern, i.e. accelerated stolonization, masculinization and the occurrence of aberrant stolons. The detailed anatomy of aberrant stolons is described. A histological study of the proventricle revealed no signs of glandular or secretory structures. The ventricle and the caeca are composed of glandular tissue but they are not involved in the reproductive and regenerative processes. CONCLUSIONS: As in other investigated syllids, the proventricle region has a significant role during stolonization and reproduction processes in Typosyllis antoni. When the proventricle region is absent, anterior and posterior regeneration are considerably deviated from the general patterns. However, proventricle ultrastructure does not show any glandular component, thereby questioning a direct involvement of this organ itself in the control of reproduction and regeneration. Our findings offer a comprehensive starting point for further studies of regeneration and reproductive control in syllids as well as annelids in general.

Syllidae mitochondrial gene order is unusually variable for Annelida.

Complete mitochondrial genomes of five syllids (Streptosyllis sp., Eusyllis blomstrandi, Myrianida brachycephala, Typosyllis antoni and Typosyllis sp.) have been obtained using Illumina sequencing. Together with two previous studied taxa (Ramisyllis multicaudata and Trypanobia cryptica), the analysed sequences represent most of the main lineages within the family Syllidae (Anoplosyllinae, Eusyllinae, Autolytinae and Syllinae). The genomic features, gene order and phylogenetic relationships are examined. Unusual for annelids, syllid mitochondrial genomes are highly variable in their gene order. Considering genomic features, such as length, skewness, gene content, and codon bias, most similar to the rest of annelids are the genomes of E. blomstrandi and M. brachycephala, while Streptosyllis sp. and the analysed sylline taxa (R. multicaudata, T. cryptica, T. antoni and Typosyllis sp.) are the most dissimilar. Two methionine tRNA's (trnM) have been found in T. antoni and Typosyllis sp. The mt genomes of these latter taxa are the longest with numerous non-coding regions. The 13 protein coding genes, as well as the rRNA's are used to perform phylogenetic analyses that recovered the relationships within the family explored before by previous authors. The gene order in Syllidae shows very different patterns. E. blomstrandi and M. prolifera show a similar pattern to the one found in Pleistoannelida; however this might have changed at least twice within Syllidae: in Streptosyllis sp. and within Syllinae. All analysed Syllinae show different gene orders, thereby illustrating more variability as all other pleistoannelids analysed so far. The information provided herein allows a more accurate reconstruction of the possible evolutionary scenarios in Syllidae.

Characterization of the complete mitochondrial genomes from Polycladida (Platyhelminthes) using next-generation sequencing.

The complete mitochondrial genomes of three polycladids, the acotylean Hoploplana elisabelloi and the cotyleans Enchiridium sp. and Prosthiostomum siphunculus have been assembled with high coverage from Illumina sequencing data. The mt genomes contain 36 genes including 12 of the 13 protein-coding genes characteristic for metazoan mitochondrial genomes, two ribosomal RNA genes, and 22 transfer RNA genes. Gene annotation, gene order, genetic code, start and stop codons and codon bias have been identified. In comparison with the well investigated parasitic Neodermata, our analysis reveals a great diversity of gene orders within Polycladida and Platyhelminthes in general. By analyzing representative genomes of the main groups of Platyhelminthes we explored the phylogenetic relationships of this group. The phylogenetic analyses strongly supported the monophyly of Polycladida, and based on a small taxon sampling suggest the monophyly of Acotylea and Cotylea.

Phylogenetic hypothesis of Sphaerodoridae Malmgren, 1867 (Annelida) and its position within Phyllodocida.

Sphaerodoridae is a small, morphologically well-defined group of annelids, with remarkable spherical tubercles covering their body surface. They have generally been considered as part of Phyllodocida, but there is no consensus about its sister group relationships. Monophyly and internal phylogenetic relationships have never been tested. We present the first phylogenetic analysis of the family, including 33 individuals of 19 species in five genera of Sphaerodoridae, and a wide representation of other Phyllodocida, based on molecular data from nuclear 18S rDNA (2240 bp), mitochondrial 16S rDNA (614 bp), and cytochrome c oxidase subunit I (657 bp). Mitochondrial and nuclear loci were analysed separately and in combination using maximum parsimony, maximum likelihood and Bayesian inference. Results show maximum support for the monophyly of Sphaerodoridae, but sister group relationships remain unclear. The genera Sphaerodoropsis and Sphaerodoridium are found to be paraphyletic. Monophyly of Clavodorum, Ephesiella and Sphaerodorum could not be tested because only one species was included for each of these genera. At least six of the morphotypes included in the study do not fit within current species descriptions, suggesting undescribed species. Finally, the transfer of Sphaerodoropsis minuta back to Sphaerodoridium is proposed.

Syllidae (Annelida: Phyllodocida) from Lizard Island, Great Barrier Reef, Australia.

Thirty species of the family Syllidae (Annelida, Phyllodocida) from Lizard Island have been identified. Three subfamilies (Eusyllinae, Exogoninae and Syllinae) are represented, as well as the currently unassigned genera Amblyosyllis and Westheidesyllis. The genus Trypanobia (Imajima & Hartman 1964), formerly considered a subgenus of Trypanosyllis, is elevated to genus rank. Seventeen species are new reports for Queensland and two are new species. Odontosyllis robustus n. sp. is characterized by a robust body and distinct colour pattern in live specimens consisting of lateral reddish-brown pigmentation on several segments, and bidentate, short and distally broad falcigers. Trypanobia cryptica n. sp. is found in association with sponges and characterized by a distinctive bright red colouration in live specimens, and one kind of simple chaeta with a short basal spur.