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Coming to terms with worms The annelids, or segmented worms, comprise one of the largest and most diverse animal phyla, found everywhere from the soil to the ocean bed. Their evolutionary relationships are poorly understood, and a reconstruction of annelid evolutionary history would be welcome. To that end, Struck et al . present a new phylogenomic analysis of 34 annelid taxa. Notable features include a division of most annelids into the Sedentaria and the Errantia, reviving a neglected 150-year-old hypothesis that the annelids developed as two major groups or clades, one specialized for a sedentary way of life and the other for a more active lifestyle. The annelids, or ringed worms, comprise one of the largest and most diverse animal phyla, and are found everywhere from garden soil to the deep sea. Their precise phylogeny has always been sketchy, but now, a new phylogenomic analysis unravels annelid evolution. Notable features include a division of most annelids into the Errantia and Sedentaria — a restitution for two groups based on classical morphology — showing how many details of anatomy and life history bear on the evolution of this important animal group. Annelida, the ringed worms, is a highly diverse animal phylum that includes more than 15,000 described species and constitutes the dominant benthic macrofauna from the intertidal zone down to the deep sea. A robust annelid phylogeny would shape our understanding of animal body-plan evolution and shed light on the bilaterian ground pattern. Traditionally, Annelida has been split into two major groups: Clitellata (earthworms and leeches) and polychaetes (bristle worms), but recent evidence suggests that other taxa that were once considered to be separate phyla (Sipuncula, Echiura and Siboglinidae (also known as Pogonophora)) should be included in Annelida 1 , 2 , 3 , 4 . However, the deep-level evolutionary relationships of Annelida are still poorly understood, and a robust reconstruction of annelid evolutionary history is needed. Here we show that phylogenomic analyses of 34 annelid taxa, using 47,953 amino acid positions, recovered a well-supported phylogeny with strong support for major splits. Our results recover chaetopterids, myzostomids and sipunculids in the basal part of the tree, although the position of Myzostomida remains uncertain owing to its long branch. The remaining taxa are split into two clades: Errantia (which includes the model annelid Platynereis ), and Sedentaria (which includes Clitellata). Ancestral character trait reconstructions indicate that these clades show adaptation to either an errant or a sedentary lifestyle, with alteration of accompanying morphological traits such as peristaltic movement, parapodia and sensory perception. Finally, life history characters in Annelida seem to be phylogenetically informative.

The simplest animal eyes are eyespots composed of two cells only: a photoreceptor and a shading pigment cell. They resemble Darwin’s ‘proto-eyes’, considered to be the first eyes to appear in animal evolution 1 , 2 , 3 , 4 . Eyespots cannot form images but enable the animal to sense the direction of light. They are characteristic for the zooplankton larvae of marine invertebrates and are thought to mediate larval swimming towards the light. Phototaxis of invertebrate larvae contributes to the vertical migration of marine plankton 5 , which is thought to represent the biggest biomass transport on Earth 6 , 7 . Yet, despite its ecological and evolutionary importance, the mechanism by which eyespots regulate phototaxis is poorly understood. Here we show how simple eyespots in marine zooplankton mediate phototactic swimming, using the marine annelid Platynereis dumerilii as a model 8 . We find that the selective illumination of one eyespot changes the beating of adjacent cilia by direct cholinergic innervation resulting in locally reduced water flow. Computer simulations of larval swimming show that these local effects are sufficient to direct the helical swimming trajectories towards the light. The computer model also shows that axial rotation of the larval body is essential for phototaxis and that helical swimming increases the precision of navigation. These results provide, to our knowledge, the first mechanistic understanding of phototaxis in a marine zooplankton larva and show how simple eyespots regulate it. We propose that the underlying direct coupling of light sensing and ciliary locomotor control was a principal feature of the proto-eye and an important landmark in the evolution of animal eyes.

Annelida is one of three animal groups possessing segmentation and is central in considerations about the evolution of different character traits. It has even been proposed that the bilaterian ancestor resembled an annelid. However, a robust phylogeny of Annelida, especially with respect to the basal relationships, has been lacking. Our study based on transcriptomic data comprising 68,750–170,497 amino acid sites from 305 to 622 proteins resolves annelid relationships, including Chaetopteridae, Amphinomidae, Sipuncula, Oweniidae, and Magelonidae in the basal part of the tree. Myzostomida, which have been indicated to belong to the basal radiation as well, are now found deeply nested within Annelida as sister group to Errantia in most analyses. On the basis of our reconstruction of a robust annelid phylogeny, we show that the basal branching taxa include a huge variety of life styles such as tube dwelling and deposit feeding, endobenthic and burrowing, tubicolous and filter feeding, and errant and carnivorous forms. Ancestral character state reconstruction suggests that the ancestral annelid possessed a pair of either sensory or grooved palps, bicellular eyes, biramous parapodia bearing simple chaeta, and lacked nuchal organs. Because the oldest fossil of Annelida is reported for Sipuncula (520 Ma), we infer that the early diversification of annelids took place at least in the Lower Cambrian.

This is the first taxonomic study of cirratulid polychaetes of the genus Kirkegaardia Blake, 2016 from Brazil. Nine new species of the genus are described from the Southern Brazilian coast (50-3000 m deep). The genus Kirkegaardia is generally subdivided into three distinct groups of species (Kirkegaardia dorsobranchialis-heterochaeta, Kirkegaardia baptisteae-tesselata and Kirkegaardia luticastella) and several out-group species for which relationships remains to be defined. In this study, new species were included in the Kirkegaardia dorsobranchialis-heterochaeta and Kirkegaardia baptisteae-tesselata groups. Kirkegaardia dorsobranchialis-heterochaeta is characterized by thoracic parapodia elevated producing a channel between the notopodia, elongate pre-setigerous region that is either entirely smooth or modified with a dorsal ridge and/or rings, and noto- and neurosetae capillaries denticulated. As belonging to this group, K. blakei sp. nov., K. brisae sp. nov., K. goytaca sp. nov., K. jongo sp. nov. and K. papaveroi sp. nov. are described here. Kirkegaardia baptisteae-tesselata includes species that lack thoracic parapodia elevated and mid-dorsal thoracic groove, although a dorsal ridge is sometimes developed. In the pre-setigerous region dorsal ridges and rings are present or absent. Most species in this group have neurosetae denticulated, and notosetae capillaries of other types. This study adds K. helenae sp. nov., K. medusa sp. nov., K. nupem sp. nov. and K. zafirae sp. nov. to the latter species group. In addition, two new records are provided for K. hampsoni. A key to cirratulid polychaete species reported from Brazilian waters is provided.

A machaeridian, Lepidocoleus hohensteini sp. nov., is described from the Hunsrueck Slate (Lower Emsian) of Germany. The available material includes a unique example preserving evidence of the soft tissues, only the second machaeridian specimen to do so and the first lepidocoleid. This specimen shows that the plates are attached to alternate segments in the trunk. The morphology is consistent with an annelid affinity of the Lepidocoleidae and confirms the unity of the Machaeridia. This discovery adds an important group to the known diversity of this famous late Palaeozoic marine Konservat-Lagerstaette.

Background Planktonic ciliated larvae are characteristic for the life cycle of marine invertebrates. Their most prominent feature is the apical organ harboring sensory cells and neurons of largely undetermined function. An elucidation of the relationships between various forms of primary larvae and apical organs is key to understanding the evolution of animal life cycles. These relationships have remained enigmatic due to the scarcity of comparative molecular data. Results To compare apical organs and larval body patterning, we have studied regionalization of the episphere, the upper hemisphere of the trochophore larva of the marine annelid Platynereis dumerilii . We examined the spatial distribution of transcription factors and of Wnt signaling components previously implicated in anterior neural development. Pharmacological activation of Wnt signaling with Gsk3β antagonists abolishes expression of apical markers, consistent with a repressive role of Wnt signaling in the specification of apical tissue. We refer to this Wnt-sensitive, six3 - and foxq2 -expressing part of the episphere as the ‘apical plate’. We also unraveled a molecular signature of the apical organ - devoid of six3 but expressing foxj , irx , nkx3 and hox - that is shared with other marine phyla including cnidarians. Finally, we characterized the cell types that form part of the apical organ by profiling by image registration, which allows parallel expression profiling of multiple cells. Besides the hox -expressing apical tuft cells, this revealed the presence of putative light- and mechanosensory as well as multiple peptidergic cell types that we compared to apical organ cell types of other animal phyla. Conclusions The similar formation of a six3+, foxq2+ apical plate, sensitive to Wnt activity and with an apical tuft in its six3- free center, is most parsimoniously explained by evolutionary conservation. We propose that a simple apical organ - comprising an apical tuft and a basal plexus innervated by sensory-neurosecretory apical plate cells - was present in the last common ancestors of cnidarians and bilaterians. One of its ancient functions would have been the control of metamorphosis. Various types of apical plate cells would then have subsequently been added to the apical organ in the divergent bilaterian lineages. Our findings support an ancient and common origin of primary ciliated larvae.

Three new species of Levinsenia were collected during a benthic survey, from 10-3,000 m deep, in Espírito Santo Basin, off the southeastern Brazilian coast. These species are L. paivai sp. nov., L. blakei sp. nov. and L. lesliae sp. nov. Members of L. paivai sp. nov. are recognized by the presence of nine pairs of well-developed and heavily ciliated branchiae, those of L. blakei sp. nov. are characterized by the presence of three pairs of small branchiae, and those of L. lesliae sp. nov., by the absence of branchiae and presence of notopodial transitional chaetae. These new species are described herein and compared to the most similar congeners. These are the first new species of Levinsenia described from off the Brazilian coast.

It has been hypothesized that a condensed nervous system with a medial ventral nerve cord is an ancestral character of Bilateria. The presence of similar dorsoventral molecular patterns along the nerve cords of vertebrates, flies, and an annelid has been interpreted as support for this scenario. Whether these similarities are generally found across the diversity of bilaterian neuroanatomies is unclear, and thus the evolutionary history of the nervous system is still contentious. Here we study representatives of Xenacoelomorpha, Rotifera, Nemertea, Brachiopoda, and Annelida to assess the conservation of the dorsoventral nerve cord patterning. None of the studied species show a conserved dorsoventral molecular regionalization of their nerve cords, not even the annelid Owenia fusiformis , whose trunk neuroanatomy parallels that of vertebrates and flies. Our findings restrict the use of molecular patterns to explain nervous system evolution, and suggest that the similarities in dorsoventral patterning and trunk neuroanatomies evolved independently in Bilateria. In bilaterian animals, the final configurations of central nervous systems seem unrelated to neuroectodermal patterning systems, so it is likely that the various architectures of the ventral nerve cords evolved convergently, many times. Convergent nervous system evolution Bilaterian animals—that is, bilaterally symmetric animals with distinct anterior and posterior ends—are often thought to have evolved from a common ancestor with a medial, ventral nerve cord. Common molecular patterns along the body axes of animals as diverse as fruit flies, annelid worms and humans support this scenario. Andreas Hejnol and colleagues look at the mediolateral neuroectodermal patterning system in a wide range of animals, including Xenoturbella (a basal bilaterian) and various lophotrochozoans (such as annelids, brachiopods and rotifers). They observe that the final anatomical configurations of the central nervous system are unrelated to the patterning system. They conclude that similar central nervous system architectures are likely to have arisen many independent times across the bilaterian group—an example of convergent evolution.

An oral tradition for microRNA Recent work suggests that microRNAs, the ubiquitous, small, non-coding genetic elements with important regulatory roles, were important in the evolution of complexity in multicellular animals. What was the role of these microRNAs when they first evolved? A deep sequencing study of the marine ragworm Platynereis dumerilii , and comparison with other bilaterian animals, suggests that the most ancient known microRNA, miR-100, was initially active in neurosecretory cells around the mouth. Other highly conserved varieties were first present in specific tissues and organ systems, such as ciliated cells and parts of the nervous system, musculature and gut. This work suggests that the last common ancestor of bilaterian animals already had all these structures. Recent work suggests that microRNAs might have been important in the evolution of complexity in multicellular animals. Here it is shown that the most ancient known microRNA, miR–100, was initially active in neurosecretory cells around the mouth. Other highly conserved varieties were first present in specific tissues and organ systems. Thus, microRNA expression was initially restricted to an ancient set of ancient animal cell types and tissues. The spectacular escalation in complexity in early bilaterian evolution correlates with a strong increase in the number of microRNAs 1 , 2 . To explore the link between the birth of ancient microRNAs and body plan evolution, we set out to determine the ancient sites of activity of conserved bilaterian microRNA families in a comparative approach. We reason that any specific localization shared between protostomes and deuterostomes (the two major superphyla of bilaterian animals) should probably reflect an ancient specificity of that microRNA in their last common ancestor. Here, we investigate the expression of conserved bilaterian microRNAs in Platynereis dumerilii , a protostome retaining ancestral bilaterian features 3 , 4 , in Capitella , another marine annelid, in the sea urchin Strongylocentrotus , a deuterostome, and in sea anemone Nematostella , representing an outgroup to the bilaterians. Our comparative data indicate that the oldest known animal microRNA, miR-100, and the related miR-125 and let-7 were initially active in neurosecretory cells located around the mouth. Other sets of ancient microRNAs were first present in locomotor ciliated cells, specific brain centres, or, more broadly, one of four major organ systems: central nervous system, sensory tissue, musculature and gut. These findings reveal that microRNA evolution and the establishment of tissue identities were closely coupled in bilaterian evolution. Also, they outline a minimum set of cell types and tissues that existed in the protostome–deuterostome ancestor.

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.