Novel molecular resources for single-specimen barcoding of enigmatic crustacean y-larvae.

Despite discovery more than 100years ago and documented global occurrence from shallow waters to the deep sea, the life cycle of the enigmatic crustacean y-larvae isincompletely understood and adult forms remain unknown. To date, only 2 of the 17 formally described species, all based on larval stages, have been investigated using an integrative taxonomic approach. This approach provided descriptions of the morphology of the naupliar and cyprid stages, and made use of exuvial voucher material and DNA barcodes. To improve our knowledge about the evolutionary history and ecological importance of y-larvae, we developed a novel protocol that maximises the amount of morpho-ecological and molecular data that can be harvested from single larval specimens. This includes single-specimen DNA barcoding and daily imaging of y-nauplii reared in culture dishes, mounting of the last naupliar exuviae on a slide as a reference voucher, live imaging of the y-cyprid instar that follows, and fixation, DNA extraction, amplification and sequencing of the y-cyprid specimen. Through development and testing of a suite of new primers for both nuclear and mitochondrial protein-coding and ribosomal genes, we showcase how new sequence data can be used to estimate the phylogeny of Facetotecta. We expect that our novel procedure will help to unravel the complex systematics of y-larvae and show how these fascinating larval forms have evolved. Moreover, we posit that our protocols should work on larval specimens from a diverse array of moulting marine invertebrate taxa.

Phylogeny and systematics of Aphroditiformia.

Aphroditiformia represents one of the most successful radiations of annelids, and is therefore an interesting model to understand morphological and functional evolution. Previous phylogenetic analyses yielded most families as monophyletic but excluded anchialine and interstitial species while failing to recover relationships within Sigalionidae. Here we address these shortcomings through the analysis of four molecular markers and 87 morphological characters sampled across 127 species under the assumptions of parsimony and model-based methods. Of the 34 newly sequenced taxa, five anchialine and 24 interstitial species were included, with increased representation of Sigalionidae. An additional 28 elusive Sigalionidae taxa were included, represented only by morphological partitions. Molecular and morphological partitions were evaluated under exhaustive sensitivity analyses, testing the effects of alignment algorithms and optimization criteria on tree topologies. Our trees congruently recovered six clades corresponding to the families within Aphroditiformia: Acoetidae, Aphroditidae, Eulepethidae, Iphionidae, Polynoidae and Sigalionidae, respectively. An anchialine polynoid lineage was nested among strictly deep sea species, and interstitial pisionids and pholoids formed two independent clades nested within Sigalionidae. Additionally, Sigalionidae resulted in four clades, defined by combinations of apomorphies, and hereby we propose the subfamilies Pelogeniinae, Pholoinae, Pisioninae, Sthenelanellinae, as well as the provisionally included polyphyletic Sigalioninae.

Description of Elisesione imajimai sp. nov. From Japan (Annelida: Hesionidae) and A Redescription of E. problematica (Wesenberg-Lund, 1950) and Its Confirmation Within Hesionini.

Naoto Jimi, Danny Eibye-Jacobsen, and Sergio I. Salazar-Vallejo (2018) Elisesione imajimai sp. nov. is newly described based on specimens collected in Sagami Bay, Japan, at a depth of 150-250 m. This new species resembles E. problematica (Wesenberg-Lund, 1950) from Iceland, but differs because in E. imajimai sp. nov. the ventral cirri surpass the tips of the neurochaetal lobes, reaching to the medial part of the neurochaetal bundle; the palps are 1/2-4/5 as long as the antennae; the aciculae are pale brownish; and the dorsal integument shows 9-10 transverse wrinkles per segment, whereas in E. problematica the ventral cirri hardly reach the tips of the neurochaetal lobes; the palps are as long as the antennae; the aciculae are black; and there are 15 transverse wrinkles per segment. A morphological redescription of E. problematica based on its type material and a revised key to identify species in the genus are also provided. A phylogenetic analysis based upon four genes (COI, 16S, 18S, 28S) confirms its position within Hesionini.

The impact of fossil data on annelid phylogeny inferred from discrete morphological characters.

As a result of their plastic body plan, the relationships of the annelid worms and even the taxonomic makeup of the phylum have long been contentious. Morphological cladistic analyses have typically recovered a monophyletic Polychaeta, with the simple-bodied forms assigned to an early-diverging clade or grade. This is in stark contrast to molecular trees, in which polychaetes are paraphyletic and include clitellates, echiurans and sipunculans. Cambrian stem group annelid body fossils are complex-bodied polychaetes that possess well-developed parapodia and paired head appendages (palps), suggesting that the root of annelids is misplaced in morphological trees. We present a reinvestigation of the morphology of key fossil taxa and include them in a comprehensive phylogenetic analysis of annelids. Analyses using probabilistic methods and both equal- and implied-weights parsimony recover paraphyletic polychaetes and support the conclusion that echiurans and clitellates are derived polychaetes. Morphological trees including fossils depict two main clades of crown-group annelids that are similar, but not identical, to Errantia and Sedentaria, the fundamental groupings in transcriptomic analyses. Removing fossils yields trees that are often less resolved and/or root the tree in greater conflict with molecular topologies. While there are many topological similarities between the analyses herein and recent phylogenomic hypotheses, differences include the exclusion of Sipuncula from Annelida and the taxa forming the deepest crown-group divergences.

Description of three new species of Eumida Malmgren, 1865 (Phyllodocidae, Annelida) from Southern and Southeastern Brazil.

We describe three new species of Eumida from shallow estuarine and shelf bottoms from south and southeastern Brazil as part of a regional survey of the family Phyllodocidae. Previous regional records of Eumida sanguinea (Örsted, 1843) are herein referred to Eumida dracodermica sp. nov. The two species differ in the size of the prostomium, the distribution of micropapillae on the proboscis, and the shape of the dorsal cirri. Eumida macrophthalma sp. nov. has much larger eyes than the other two species described herein. The dorsal cirri on anterior median segments are rounded and on median and posterior segments they are asymmetrical and cordiform, longer than the dorsal cirri in Eumida dracodermica sp. nov. The lanceolate median dorsal cirri in Eumida delicata sp. nov. are only shared with E. ockelmanni Eibye-Jacobsen, 1987. It is also possible to diagnose the three species by their dorsal whitish pigmentation patterns on the first segments of living or well conserved animals.

Annulata örstediana: publication dates, composition and annotated taxonomic list, with some comments on Hemipodus (Polychaeta: Glyceridae).

The collecting trips by Ørsted and Kröyer in Central and South America resulted in a series of papers by Grube, the Annulata örstediana which contained the proposal for seven genera and descriptions for 84 species. There are some problems dealing with the correct number of contributions, their publication dates, the correct citation for the authors for each species name and sometimes even for the type locality. In order to improve the current situation, we have consulted the original publications together with the corresponding collections. Our results indicate that there were only three parts in the series which were published in the journal, and repagination in reprints sometimes caused some further confusion. Additionally, we provided some review comments on the species current status and we added a short nomenclatural note on Hemipodus (Polychaeta: Glyceridae) trying to clarify the correct generic name and type species.

Annulata örstediana: publication dates, composition and annotated taxonomic list, with some comments on Hemipodus (Polychaeta: Glyceridae)

The collecting trips by Ørsted and Kröyer in Central and South America resulted in a series of papers by Grube, the Annulata örstediana which contained the proposal for seven genera and descriptions for 84 species. There are some problems dealing with the correct number of contributions, their publication dates, the correct citation for the authors for each species name and sometimes even for the type locality. In order to improve the current situation, we have consulted the original publications together with the corresponding collections. Our results indicate that there were only three parts in the series which were published in the journal, and repagination in reprints sometimes caused some further confusion. Additionally, we provided some review comments on the species current status and we added a short nomenclatural note on Hemipodus (Polychaeta: Glyceridae) trying to clarify the correct generic name and type species.

Los viajes de recolecta por Ørsted y Kröyer en Centroamérica y Sudamérica resultaron en una serie de notas por Grube, los Annulata örstediana que contienen la propuesta de siete géneros y 84 especies. Hay algunos problemas en relación con el número correcto de contribuciones, sus fechas de publicación, la citación correcta de los autores para cada nombre específico y a veces incluso para la localidad tipo. Para mejorar la situación, hemos consultado las publicaciones originales junto con las colecciones correspondientes. Nuestros resultados indican que hubo tres partes en la serie, que fueron publicadas en la revista y cuyas páginas fueron re-enumeradas para las separatas lo que a veces ocasionó confusiones adicionales. Además, brindamos algunos comentarios para revisar el estado de cada especie y agregamos una breve nota nomenclatural sobre Hemipodus (Polychaeta: Glyceridae) para aclarar el nombre correcto del género y la especie tipo.

An Early Cambrian stem polychaete with pygidial cirri.

The oldest annelid fossils are polychaetes from the Cambrian Period. They are representatives of the annelid stem group and thus vital in any discussion of how we polarize the evolution of the crown group. Here, we describe a fossil polychaete from the Early Cambrian Sirius Passet fauna, Pygocirrus butyricampum gen. et sp. nov., with structures identified as pygidial cirri, which are recorded for the first time from Cambrian annelids. The body is slender and has biramous parapodia with chaetae organized in laterally oriented bundles. The presence of pygidial cirri is one of the characters that hitherto has defined the annelid crown group, which diversified during the Cambrian-Ordovician transition. The newly described fossil shows that this character had already developed within the total group by the Early Cambrian.

Jaw morphology and ontogeny in five species of Ophryotrocha.

Detailed scanning electron microscopy of jaws within the genus Ophryotrocha (Dorvilleidae, Annelida) was performed on 871 jaw parts. The investigations resulted in new understandings of the ontogeny and jaw morphology and have systematic implications for the family. Five species in the genus (Ophryotrocha alborana, O. diadema, O. gracilis, O. hartmanni, and O. labronica pacifica) were kept in culture, and the development of the jaws was studied by sampling throughout their life history. Ophryotrocha species have mandibular plates that remain the same throughout ontogeny, whereas the posterior shafts elongate. Both mandibular plate morphology and shaft ontogeny have species-specific distinctions. In Ophryotrocha, the maxillae can be assigned to three to four distinct types, which are replaced by moulting. The maxillary morphology and developmental stages at which moults occur are species specific, although with broad intervals. A redefinition is given for some of the basic jaw elements, and new homologies are proposed for structures that are also present across other dorvilleid taxa.

Ultrastructure of the extensively developed nuchal organs of Laonice bahusiensis (Annelida: Canalipalpata: Spionidae).

The nuchal organs of annelid Laonice bahusiensis (Spionidae) from northern Europe have been studied using scanning and transmission electron microscopy. L. bahusiensis is the first spionid species in which extensively developed, continuous nuchal organs are described. The nuchal organs of this genus are the longest known among polychaete annelids. They consist of paired double bands extending from the prostomium on a mid-dorsal caruncle for about 24-30 setigers. Their microanatomy corresponds to the general structural plan of nuchal organs: there are ciliated supporting cells and bipolar sensory cells with sensory cilia traversing an olfactory chamber. The organs are overlaid by a secondary paving-stone-like cover and innervated by one pair of longitudinally elongated nuchal nerves. These findings clearly favor the hypothesis that the paired, extensively developed ciliated structures found in some Spionidae are homologous with the prostomial nuchal organs characteristic of polychaete annelids.