Search

AbstractWe present the mitochondrial genome of the deep-sea, epibenthic, irregular echinoid Echinocrepis rostrata, representing the first sequenced mitogenome of the order Holasteroida. The length of the complete E. rostrata mitochondrial genome is 15,716 base pairs, and its GC content is 34.87%. It contains 13 protein-coding genes, two rRNA genes, and 22 tRNA genes, whose order is identical to that of all other available echinoid mitogenomes. Phylogenetic analysis of available mitochondrial genomes, based on all coding loci, places E. rostrata as the sister group to spatangoids (heart urchins).

Abstract Background Common seadragons (Phyllopteryx taeniolatus, Syngnathidae) are an emblem of the diverse endemic fauna of Australia’s southern rocky reefs, the newly recognized “Great Southern Reef.” A lack of assessments spanning this global biodiversity hotspot in its entirety is currently hampering an understanding of the factors that have contributed to its diversity. The common seadragon has a wide range across Australia's entire temperate south and includes a geogenetic break over a former land bridge, which has called its status as a single species into question. As a popular aquarium display that sells for high prices, common seadragons are also vulnerable to illegal capture. Results Here, we provide range-wide nuclear sequences (986 variable Ultraconserved Elements) for 198 individuals and mitochondrial genomes for 140 individuals to assess species status, identify genetic units and their diversity, and trace the source of two poached individuals. Using published data of the other two seadragon species, we found that lineages of common seadragons have diverged relatively recently (

Abstract Bacterial symbioses allow annelids to colonise extreme ecological niches, such as hydrothermal vents and whale falls. Yet, the genetic principles sustaining these symbioses remain unclear. Here, we show that different genomic adaptations underpin the symbioses of phylogenetically related annelids with distinct nutritional strategies. Genome compaction and extensive gene losses distinguish the heterotrophic symbiosis of the bone-eating worm Osedax frankpressi from the chemoautotrophic symbiosis of deep-sea Vestimentifera. Osedax’s endosymbionts complement many of the host’s metabolic deficiencies, including the loss of pathways to recycle nitrogen and synthesise some amino acids. Osedax’s endosymbionts possess the glyoxylate cycle, which could allow more efficient catabolism of bone-derived nutrients and the production of carbohydrates from fatty acids. Unlike in most Vestimentifera, innate immunity genes are reduced in O. frankpressi, which, however, has an expansion of matrix metalloproteases to digest collagen. Our study supports that distinct nutritional interactions influence host genome evolution differently in highly specialised symbioses.

Spirorbinae, a ubiquitous group of marine calcareous tubeworms with a small body size as adults, have a fascinating diversity of brooding modes that form the basis for their taxonomic division into six tribes (traditionally subfamilies): in-tube incubation, with varying degrees of attachment to adult structures (four tribes), and external incubation in a modified radiole (opercular brood chambers; two tribes). We investigated the evolutionary transitions among these brooding modes. Phylogenetic reconstruction with molecular (28s and 18s rDNA) and morphological data (83 characters) among 36 taxa (32 ingroup spirorbins; 4 filogranin outgroups) of the combined data set, using maximum parsimony, maximum likelihood, and Bayesian analyses, inferred Spirorbinae to be monophyletic, with strong support for the monophyly for five tribes (Circeini, Januini, Romanchellini, Paralaeospirini and Spirorbini), but non-monophyly for Pileolariini. However, deeper relationships among some tribes remain unresolved. Neomicrorbis was found to be the sistergroup to all other Spirorbinae. Alternative coding strategies for assessing the ancestral state reconstruction for the reproductive mode allowed for a range of conclusions as to the evolution of tube and opercular brooding in Spirorbinae. Two of the transformations suggest that opercular brooding may be ancestral for Spirorbinae, and the tube-incubating tribes may have been derived independently from opercular-brooding ancestors.

Two new species of Hesionidae, Parahesione pulvinata sp. nov. and Parahesione apiculata sp. nov. are described based on materials collected at tidal flats in Okinawa (Japan) from burrows of the ghost shrimps Neocallichirus jousseaumei and Glypturus armatus. The two new species are characterized by having eight enlarged cirri, dorsal cirrophores with dorsal foliose lobe and biramous parapodia, and by lacking median antenna. Parahesione apiculata sp. nov. has digitate lobes on the posterior margin of the dorsal foliose lobe (absent in P. pulvinata sp. nov.). The two new species were never found outside the ghost shrimp burrows, suggesting they are obligate symbionts. Phylogenetic analyses based on four concatenated genes suggest that the symbiotic lifestyle has evolved several times in Hesionidae.

ABSTRACT Osedax, the deep-sea annelid found at sunken whalefalls, is known to host Oceanospirillales bacterial endosymbionts intracellularly in specialized roots, which help it feed exclusively on vertebrate bones. Past studies, however, have also made mention of external bacteria on their trunks. During a 14-yr study, we reveal a dynamic, yet persistent, shift of Campylobacterales integrated into the epidermis of Osedax, which change over time as the whale carcass degrades on the sea floor. The Campylobacterales associated with seven species of Osedax, which comprise 67% of the bacterial community on the trunk, appear initially dominated by the genus Arcobacter (at early time points 140 mo) of whale carcass decomposition. Metagenome analysis of the epibiont metabolic capabilities suggests potential for a transition from heterotrophy to autotrophy and differences in their capacity to metabolize oxygen, carbon, nitrogen, and sulfur. Compared to free-living relatives, the Osedax epibiont genomes were enriched in transposable elements, implicating genetic exchange on the host surface, and contained numerous secretions systems with eukaryotic-like protein (ELP) domains, suggesting a long evolutionary history with these enigmatic, yet widely distributed deep-sea worms. IMPORTANCE Symbiotic associations are widespread in nature and we can expect to find them in every type of ecological niche. In the last twenty years, the myriad of functions, interactions and species comprising microbe-host associations has fueled a surge of interest and appreciation for symbiosis. During this 14-year study, we reveal a dynamic population of bacterial epibionts, integrated into the epidermis of 7 species of a deep-sea worm group that feeds exclusively on the remains of marine mammals. The bacterial genomes provide clues of a long evolutionary history with these enigmatic worms. On the host surface, they exchange genes and appear to undergo ecological succession, as the whale carcass habitat degrades over time, similar to what is observed for some free-living communities. These, and other annelid worms are important keystone species for diverse deep-sea environments, yet the role of attached external bacteria in supporting host health has received relatively little attention.

Pilargidae is a family of free-living and burrowing marine annelids. A lack of available molecular data for most of these species has precluded a molecular assessment of their phylogenetic relationships and has left uncertain the placement of Antonbruunia, which is hypothesized to be either a member of Pilargidae or its sister clade, the monotypic family Antonbruunidae. In this study, we describe the new species Antonbruunia milenae sp. nov., found at 845 m of depth off the coast of San Diego, California, USA, and we address the phylogeny of these organisms using 15 novel mitogenomes and multiple Sanger-sequenced loci. Our results show that Antonbruunia falls within Pilargidae, making Antonbruunidae a junior synonym of Pilargidae. Glyphohesione was transferred from Pilarginae to Synelminae, the previously unassigned genera Otopsis and Antonbruunia were shown to belong within Synelminae, and Hermundura was assigned to Phyllodocida incertae sedis. Sigambra was found to be non-monophyletic. Four different mitogenome gene orders were found among Pilargidae. Changes between the gene orders and the ancestral state gene order of the family were inferred. Two species have introns within the COI gene. These efforts represent a significant expansion of the available molecular resources for pilargids, as well as the basis for a more stable taxonomy.

Dorvilleids belonging to Ophryotrocha Claparède & Mecznikow, 1869 are known from deep-sea hydrothermal vents in the Pacific, Atlantic, Indian and Southern Oceans. However, how they colonized and diversified in these ecosystems has not been assessed in detail. Here, a collection of Pacific hydrothermal vent Ophryotrocha was examined using morphology and DNA markers (COI, 16S and H3). Five new species were revealed, largely expanding the diversity of the group at this habitat type. They are Ophryotrocha charlottae sp. nov., O. kailae sp. nov., O. marinae sp. nov., O. pruittae sp. nov. from eastern Pacific, and O. bohnorum sp. nov. from the western Pacific. Phylogenetic analyses based on the concatenated alignments of all three genes suggest vent habitants have been colonized several times independently within Ophryotrocha. One clade of six vent species was recovered, indicative of diversification following a colonization of hydrothermal vents, likely in the eastern Pacific. An Indian Ocean species, O. jiaolongi, was nested inside this clade and was closely related to one of the new species from the Gulf of California, diverging from it by less than 4% on COI.

Xyloplax is a genus of three species of sea stars previously found only on sunken wood in the deep ocean. Their circular and petaloid bodies, which lend them their common name “sea daisy”, and their presumed exclusive diet of wood make them an unusual and rare element of deep-sea ecosystems. We describe here the fourth species of Xyloplax from the eastern Pacific Ocean, Xyloplax princealberti n. sp., which ranges from offshore Canada to the Gulf of California (Mexico) and Costa Rica. Though sampled geographically close to another described species of Xyloplax from the northeastern Pacific, X. janetae, this new species is unique morphologically and according to available DNA data. The short abactinal spines are the most obvious feature that distinguishes X. princealberti n. sp. from other Xyloplax. The minimum distance for mitochondrial cytochrome c oxidase subunit I from Xyloplax princealberti n. sp. to the only other available Xyloplax, X. janetae, was 13.5%. We also describe Ridgeia vestimentiferan tubeworm bushes from active hydrothermal vents as a new Xyloplax habitat, the first record of a non-wood substrate, and a new reproductive strategy, simultaneous hermaphroditism, for this genus. We generated the first mitochondrial genome for a member of Xyloplax and analyzed it with other available asteroid data using nucleotide-coding or amino acid (for protein-coding genes) plus nucleotide coding (for rRNA genes). The nucleotide-coding results place Xylopax as part of the clade Velatida, consistent with a previous phylogenomic analysis that included Xyloplax princealberti n. sp. (as Xyloplax sp.), though the placement of Velatida within Asteroidea differed. The amino acid plus nucleotide coding recovered Velatida to be a grade with X. princealberti n. sp. as sister group to all other Asteroidea.

Abstract Background Seahorses, seadragons, pygmy pipehorses, and pipefishes (Syngnathidae, Syngnathiformes) are among the most recognizable groups of fishes because of their derived morphology, unusual life history, and worldwide distribution. Despite previous phylogenetic studies and recent new species descriptions of syngnathids, the evolutionary relationships among several major groups within this family remain unresolved. Results Here, we provide a reconstruction of syngnathid phylogeny based on genome-wide sampling of 1314 ultraconserved elements (UCEs) and expanded taxon sampling to assess the current taxonomy and as a basis for macroevolutionary insights. We sequenced a total of 244 new specimens across 117 species and combined with published UCE data for a total of 183 species of Syngnathidae, about 62% of the described species diversity, to compile the most data-rich phylogeny to date. We estimated divergence times using 14 syngnathiform fossils, including nine fossils with newly proposed phylogenetic affinities, to better characterize current and historical biogeographical patterns, and to reconstruct diversification through time. We present a phylogenetic hypothesis that is well-supported and provides several notable insights into syngnathid evolution. We found nine non-monophyletic genera, evidence for seven cryptic species, five potentially invalid synonyms, and identified a novel sister group to the seahorses, the Indo-Pacific pipefishes Halicampus macrorhynchus and H. punctatus. In addition, the morphologically distinct southwest Pacific seahorse Hippocampus jugumus was recovered as the sister to all other non-pygmy seahorses. As found in many other groups, a high proportion of syngnathid lineages appear to have originated in the Central Indo-Pacific and subsequently dispersed to adjoining regions. Conversely, we also found an unusually high subsequent return of lineages from southern Australasia to the Central Indo-Pacific. Diversification rates rose abruptly during the Middle Miocene Climate Transition and peaked after the closure of the Tethys Sea. Conclusions Our results reveal a previously underappreciated diversity of syngnathid lineages. The observed biogeographic patterns suggest a significant role of the southern Australasian region as a source and sink of lineages. Shifts in diversification rates imply possible links to declining global temperatures, the separation of the Atlantic and Pacific faunas, and the environmental changes associated with these events.

Here, we present the first mitochondrial genome of Eiconaxius baja. The mitogenome contains 13 protein-coding genes (PCGs), two rRNA genes, and 22 tRNA genes. The total length of the complete E. baja mitochondrial genome is 16,212 base pairs, and the GC content is 26.82%. The gene order is consistent with that of Eiconaxius serratus, and most other members of Axiidea. Phylogenetic analysis based on 13 PCGs places E. baja sister to E. serratus within Axiidae.

Abstract The North Pacific bigeye octopus, Octopus californicus (Berry, 1911) is a cold‐water, deep‐sea octopod. Little is known about their biology due to difficulty accessing their natural habitat and obtaining live specimens. Although they are a frequent bycatch product in commercial bottom trawl fisheries, individuals of this species have rarely been raised in captivity and their embryonic development has not yet been documented. Considering these limitations, we were fortunate to have witnessed this process leading to successful hatching in an aquarium setting. Here, we present a brief observational account of the first‐known record of brooding, development, and hatching of fertilized eggs for O. californicus. The incubation time was a maximum of 10 months at a temperature between 8–10°C and embryos hatched over a period of 2.5 months. While more detailed research is needed, this preliminary information contributes to our limited knowledge of this species and supports life history theories of prolonged embryonic development under colder temperatures.

The taxonomy of ribbon worms (Nemertea) is particularly challenging due to the sparsity of distinct morphological characters, causing a significant underestimation of the group’s true diversity. The number of named deep-sea species is very limited and there is a vast number of undescribed deep-sea nemerteans still to be discovered. In this paper we figuratively ‘cut the ribbon’ and name seven new species of monostiliferous hoplonemerteans from seeps and seamounts along the Costa Rican margin, one from seeps along the Oregon margin, and one from vents of the Juan de Fuca Ridge, USA. The species Chernyshevia escarpiaphila gen. et sp. nov. and five species of the genus Alvinonemertes gen. nov. (Alvinonemertes dariae gen. et sp. nov., Alvinonemertes dagmarae gen. et sp. nov., Alvinonemertes christianeae gen. et sp. nov., Alvinonemertes claudiae gen. et sp. nov., Alvinonemertes tatjanae gen. et sp. nov.) represent Oerstediina, whereas the three species Tetrastemma sundbergi sp. nov., Tetrastemma polyakovae sp. nov., and Tetrastemma strandae sp. nov. represent Amphiporina. One species of tubulanid palaeonemerteans is described but not provided with a species name due to lacking sequence data for comparison. Additionally, we provide sequence data for one lineid heteronemertean, one reptant hoplonemertean, and two further eumonostiliferan hoplonemertean species. We use an integrative, turbotaxonomic approach combining DNA sequence data with concise morphological descriptions and fully digitized serial histological sections made available as cybertypes.

Abstract Background Numerous deep-sea invertebrates, at both hydrothermal vents and methane seeps, have formed symbiotic associations with internal chemosynthetic bacteria in order to harness inorganic energy sources typically unavailable to animals. Despite success in nearly all marine habitats and their well-known associations with photosynthetic symbionts, Cnidaria remain one of the only phyla present in the deep-sea without a clearly documented example of dependence on chemosynthetic symbionts. Results A new chemosynthetic symbiosis between the sea anemone Ostiactis pearseae and intracellular bacteria was discovered at ~ 3700 m deep hydrothermal vents in the southern Pescadero Basin, Gulf of California. Unlike most sea anemones observed from chemically reduced habitats, this species was observed in and amongst vigorously venting fluids, side-by-side with the chemosynthetic tubeworm Oasisia aff. alvinae. Individuals of O. pearseae displayed carbon, nitrogen, and sulfur tissue isotope values suggestive of a nutritional strategy distinct from the suspension feeding or prey capture conventionally employed by sea anemones. Molecular and microscopic evidence confirmed the presence of intracellular SUP05-related bacteria housed in the tentacle epidermis of O. pearseae specimens collected from 5 hydrothermally active structures within two vent fields ~ 2 km apart. SUP05 bacteria (Thioglobaceae) dominated the O. pearseae bacterial community, but were not recovered from other nearby anemones, and were generally rare in the surrounding water. Further, the specific Ostiactis-associated SUP05 phylotypes were not detected in the environment, indicating a specific association. Two unusual candidate bacterial phyla (the OD1 and BD1-5 groups) appear to associate exclusively with O. pearseae and may play a role in symbiont sulfur cycling. Conclusion The Cnidarian Ostiactis pearseae maintains a physical and nutritional alliance with chemosynthetic bacteria. The mixotrophic nature of this symbiosis is consistent with what is known about other cnidarians and the SUP05 bacterial group, in that they both form dynamic relationships to succeed in nature. The advantages gained by appropriating metabolic and structural resources from each other presumably contribute to their striking abundance in the Pescadero Basin, at the deepest known hydrothermal vents in the Pacific Ocean.

Polynoidae Kinberg, 1856 has five branchiate genera: Branchipolynoe Pettibone, 1984, Branchinotogluma Pettibone, 1985, Branchiplicatus Pettibone, 1985, Peinaleopolynoe Desbruyères & Laubier, 1988, and Thermopolynoe Miura, 1994, all native to deep-sea, chemosynthetic-based habitats. Of these, Peinaleopolynoe has two accepted species; Peinaleopolynoe sillardi Desbruyères & Laubier, 1988 (Atlantic Ocean) and Peinaleopolynoe santacatalina Pettibone, 1993 (East Pacific Ocean). The goal of this study was to assess the phylogenetic position of Peinaleopolynoe, utilizing DNA sequences from a broad sampling of deep-sea polynoids. Representatives from all five branchiate genera were included, several species of which were sampled from near the type localities; Branchinotogluma sandersi Pettibone, 1985 from the Galápagos Rift (E/V “Nautilus”); Peinaleopolynoe sillardi from organic remains in the Atlantic Ocean; Peinaleopolynoe santacatalina from a whalefall off southern California (R/V “Western Flyer”) and Thermopolynoe branchiata Miura, 1994 from Lau Back-Arc Basin in the western Pacific (R/V “Melville”). Phylogenetic analyses were conducted using mitochondrial (COI, 16S rRNA, and CytB) and nuclear (18S rRNA, 28S rRNA, and H3) genes. The analyses revealed four new Peinaleopolynoe species from the Pacific Ocean that are formally described here: Peinaleopolynoe orphanae Hatch & Rouse, sp. nov., type locality Pescadero Basin in the Gulf of California, Mexico (R/V “Western Flyer”); Peinaleopolynoe elvisi Hatch & Rouse, sp. nov. and Peinaleopolynoe goffrediae Hatch & Rouse, sp. nov., both with a type locality in Monterey Canyon off California (R/V “Western Flyer”) and Peinaleopolynoe mineoi Hatch & Rouse, sp. nov. from Costa Rica methane seeps (R/V “Falkor”). In addition to DNA sequence data, the monophyly of Peinaleopolynoe is supported by the presence of ventral papillae on segments 12–15. The results also demonstrated the paraphyly of Branchinotogluma and Lepidonotopodium Pettibone, 1983 and taxonomic revision of these genera is required. We apply the subfamily name Lepidonotopodinae Pettibone 1983, for the clade comprised of Branchipolynoe, Branchinotogluma, Bathykurila, Branchiplicatus, Lepidonotopodium, Levensteiniella Pettibone, 1985, Thermopolynoe, and Peinaleopolynoe.

Serpulidae Rafinesque, 1815 is a speciose group of polychaetes that all inhabit calcareous tubes. The family was traditionally subdivided into Serpulinae, Filograninae, and Spirorbinae. Recent phylogenetic analyses have suggested that both Filograninae and Serpulinae are paraphyletic, though with limited sampling. Here we report the first phylogenetic analysis of Serpulidae based on comprehensive sampling of genera (though excluding most spirorbin genera). We include a much-needed revision of serpulid taxonomy based on a phylogenetic hypothesis derived from both morphological and molecular data. We analysed 18S, 28S, histone H3 ribosomal nuclear DNA and cytochrome b (cytb) mitochondrial sequences, combined with morphological data. The proposed new classification includes the re-formulated Serpulinae (with tribes Serpulini and Ficopomatini), Spirorbinae, and Filograninae, with apomorphies highlighted for major taxa.

Here, we report the 15,103 bp mitochondrial genome of the freshwater fabriciid tubeworm Manayunkia occidentalis. We recovered 13 protein-coding genes, 2 rRNA, and 22 tRNA. The gene order is consistent with the conserved pattern observed in most annelids.

Abstract Background Marine invertebrates are abundant and diverse on the continental shelf in Antarctica, but little is known about their parasitic counterparts. Endoparasites are especially understudied because they often possess highly modified body plans that pose problems for their identification. Asterophila, a genus of endoparasitic gastropod in the family Eulimidae, forms cysts in the arms and central discs of asteroid sea stars. There are currently four known species in this genus, one of which has been described from the Antarctic Peninsula (A. perknasteri). This study employs molecular and morphological data to investigate the diversity of Asterophila in Antarctica and explore cophylogenetic patterns between host and parasite. Results A maximum-likelihood phylogeny of Asterophila and subsequent species-delimitation analysis uncovered nine well-supported putative species, eight of which are new to science. Most Asterophila species were found on a single host species, but four species were found on multiple hosts from one or two closely related genera, showing phylogenetic conservatism of host use. Both distance-based and event-based cophylogenetic analyses uncovered a strong signal of coevolution in this system, but most associations were explained by non-cospeciation events. Discussion The prevalence of duplication and host-switching events in Asterophila and its asteroid hosts suggests that synchronous evolution may be rare even in obligate endoparasitic systems. The apparent restricted distribution of Asterophila from around the Scotia Arc may be an artefact of concentrated sampling in the area and a low obvious prevalence of infection. Given the richness of parasites on a global scale, their role in promoting host diversification, and the threat of their loss through coextinction, future work should continue to investigate parasite diversity and coevolution in vulnerable ecosystems.

Wood boring bivalves of the family Xylophagaidae inhabit sunken wood on the deep-sea floor where they play a key role in the degradation of this organic matter in the ocean. The patchiness of wood-fall habitats is impeding targeted sampling and little is therefore known on xylophagaid biology. We investigated for the first time the diversity and biogeography of Xylophagaidae in the NE-Atlantic and the Mediterranean over a broad geographic range and in various water depths using experimental wood deployments. We combined morphological and molecular analyses for species discrimination. A phylogenetic reconstruction based on 18S and 28S rRNA and COI genes revealed non-monophyly of the type genus, XylophagaTurton (1822), and led us to revise the taxonomy and erect the genus Xylonora gen. nov. COI haplotypes of the most abundant species revealed broad Atlanto-Mediterranean genetic connectivity for Xylophaga dorsalis and Xylonora atlantica new comb., while genetic connectivity appears limited for Abditoconus brava across the entrance of the Mediterranean. We provide the first COI barcode data for Xylophagaidae as a solid base for future taxonomic work. Wood deployments in a broad geographic range provided a powerful tool for research on Xylophagaidae allowing for conclusions on ecological requirements of xylophagaid species.

We describe a long-unnamed Chaetopterus Cuvier, 1830 species from southern California, using a combination of DNA barcoding and detailed morphological investigation employing high-resolution X-ray microtomography (micro-CT). Chaetopterus dewysee sp. nov. is not only one of the most dominant annelids in the benthic communities of the shallow end of the La Jolla submarine canyon, but also a well-established model for studying bioluminescence and has a published transcriptome. The description and naming of this southern Californian Chaetopterus is a step towards the much-needed revision of the group’s taxonomy and towards resolving the confusion over the ʻcosmopolitanʼ Chaetopterus variopedatus species complex. Micro-CT data showing details of both internal and external anatomy has been made freely available as the first annelid cybertype.

Abstract Background Echinoidea is a clade of marine animals including sea urchins, heart urchins, sand dollars and sea biscuits. Found in benthic habitats across all latitudes, echinoids are key components of marine communities such as coral reefs and kelp forests. A little over 1000 species inhabit the oceans today, a diversity that traces its roots back at least to the Permian. Although much effort has been devoted to elucidating the echinoid tree of life using a variety of morphological data, molecular attempts have relied on only a handful of genes. Both of these approaches have had limited success at resolving the deepest nodes of the tree, and their disagreement over the positions of a number of clades remains unresolved. Results We performed de novo sequencing and assembly of 17 transcriptomes to complement available genomic resources of sea urchins and produce the first phylogenomic analysis of the clade. Multiple methods of probabilistic inference recovered identical topologies, with virtually all nodes showing maximum support. In contrast, the coalescent-based method ASTRAL-II resolved one node differently, a result apparently driven by gene tree error induced by evolutionary rate heterogeneity. Regardless of the method employed, our phylogenetic structure deviates from the currently accepted classification of echinoids, with neither Acroechinoidea (all euechinoids except echinothurioids), nor Clypeasteroida (sand dollars and sea biscuits) being monophyletic as currently defined. We show that phylogenetic signal for novel resolutions of these lineages is strong and distributed throughout the genome, and fail to recover systematic biases as drivers of our results. Conclusions Our investigation substantially augments the molecular resources available for sea urchins, providing the first transcriptomes for many of its main lineages. Using this expanded genomic dataset, we resolve the position of several clades in agreement with early molecular analyses but in disagreement with morphological data. Our efforts settle multiple phylogenetic uncertainties, including the position of the enigmatic deep-sea echinothurioids and the identity of the sister clade to sand dollars. We offer a detailed assessment of evolutionary scenarios that could reconcile our findings with morphological evidence, opening up new lines of research into the development and evolutionary history of this ancient clade.

The scale-worm family Iphionidae consists of four genera. Of these, Thermiphione has two accepted species, both native to hydrothermal vents in the Pacific Ocean; T. fijiensis Miura, 1994 (West Pacific) and T. tufari Hartmann-Schröder, 1992 (East Pacific Rise). Iphionella is also known from the Pacific, and has two recognized species; Iphionella risensis Pettibone, 1986 (East Pacific Rise, hydrothermal vents) and I. philippinensis Pettibone, 1986 (West Pacific, deep sea). In this study, phylogenetic analyses of Iphionidae from various hydrothermal vent systems of the Pacific Ocean were conducted utilizing morphology and mitochondrial (COI and 16S rRNA) and nuclear (18S and 28S rRNA) genes. The results revealed a new iphionid species, described here as Thermiphione rapanui sp. n. The analyses also demonstrated the paraphyly of Thermiphione, requiring Iphionella risensis to be referred to the genus, as Thermiphione risensis (Pettibone, 1986).

Stomatopoda, commonly known as mantis shrimps, are notable for their enlarged second maxillipeds encompassing the raptorial claw. The form of the claw can be used to divide them into two basic groups: smashers and spearers. Previous phylogenetic studies of Stomatopoda have focused on morphology or a few genes, though there have been whole mitochondrial genomes published for 15 members of Stomatopoda. However, the sampling has been somewhat limited with key taxa not included. Here, nine additional stomatopod mitochondrial genomes were generated and combined with the other available mitogenomes for a phylogenetic analysis. We used the 13 protein coding genes, as well as 12S rRNA, 16S rRNA genes, and included nuclear 18S rRNA gene sequences. Different rooting options were used for the analyses: (1) single and multiple outgroups from various eumalocostracan relatives and (2) a stomatopod-only dataset, with Hemisquilla californiensis used to root the topologies, based on the current hypothesis that Hemisquilla is the sister group to the rest of Stomatopoda. The eumalocostracan-rooted analyses all showed H. californiensis nested within Stomatopoda, raising doubts as to previous hypotheses as to its placement. Allowing for the rooting difference, the H. californiensis outgroup datasets had the same tree topology as the eumalocostracan outgroup datasets with slight variation at poorly supported nodes. Of the major taxonomic groupings sampled to date, Squilloidea was generally found to be monophyletic while Gonodactyloidea was not. The position of H. californiensis was found inside its superfamily, Gonodactyloidea, and grouped in a weakly supported clade containing Odontodactylus havanensis and Lysiosquillina maculata for the eumalocostracan-rooted datasets. An ancestral state reconstruction was performed on the raptorial claw form and provides support that spearing is the ancestral state for extant Stomatopoda, with smashing evolving subsequently one or more times.

Abstract Background A range of higher animal taxa are shared across various chemosynthesis-based ecosystems (CBEs), which demonstrates the evolutionary link between these habitats, but on a global scale the number of species inhabiting multiple CBEs is low. The factors shaping the distributions and habitat specificity of animals within CBEs are poorly understood, but geographic proximity of habitats, depth and substratum have been suggested as important. Biogeographic studies have indicated that intermediate habitats such as sedimented vents play an important part in the diversification of taxa within CBEs, but this has not been assessed in a phylogenetic framework. Ampharetid annelids are one of the most commonly encountered animal groups in CBEs, making them a good model taxon to study the evolution of habitat use in heterotrophic animals. Here we present a review of the habitat use of ampharetid species in CBEs, and a multi-gene phylogeny of Ampharetidae, with increased taxon sampling compared to previous studies. Results The review of microhabitats showed that many ampharetid species have a wide niche in terms of temperature and substratum. Depth may be limiting some species to a certain habitat, and trophic ecology and/or competition are identified as other potentially relevant factors. The phylogeny revealed that ampharetids have adapted into CBEs at least four times independently, with subsequent diversification, and shifts between ecosystems have happened in each of these clades. Evolutionary transitions are found to occur both from seep to vent and vent to seep, and the results indicate a role of sedimented vents in the transition between bare-rock vents and seeps. Conclusion The high number of ampharetid species recently described from CBEs, and the putative new species included in the present phylogeny, indicates that there is considerable diversity still to be discovered. This study provides a molecular framework for future studies to build upon and identifies some ecological and evolutionary hypotheses to be tested as new data is produced.

Abstract Bacteria represent the most genetically diverse kingdom of life. While great progress has been made in describing this diversity, it remains difficult to identify the phylogenetic and ecological characteristics that delineate groups of bacteria that possess species-like properties. One major challenge associated with species delineations is that not all shared genes have the same evolutionary history, and thus the choice of loci can have a major impact on phylogenetic reconstruction. Sequencing the genomes of large numbers of closely related strains provides new opportunities to distinguish ancestral from acquired alleles and assess the effects of recombination on phylogenetic inference. Here we analyzed the genomes of 119 strains of the marine actinomycete genus Salinispora, which is currently comprised of three named species that share 99% 16S rRNA gene sequence identity. While 63% of the core genome showed evidence of recombination, this had no effect on species-level phylogenomic resolution. Recombination did however blur intra-species relationships and biogeographic resolution. The genome-wide average nucleotide identity provided a new perspective on Salinispora diversity, revealing as many as seven new species. Patterns of orthologous group distributions reveal a genetic basis to delineation the candidate taxa and insight into the levels of genetic cohesion associated with bacterial species.

Terebelliformia—“spaghetti worms” and their allies—are speciose and ubiquitous marine annelids but our understanding of how their morphological and ecological diversity evolved is hampered by an uncertain delineation of lineages and their phylogenetic relationships. Here, we analyzed transcriptomes of 20 terebelliforms and an outgroup to build a robust phylogeny of the main lineages grounded on 12,674 orthologous genes. We then supplemented this backbone phylogeny with a denser sampling of 121 species using five genes and 90 morphological characters to elucidate fine-scale relationships. The monophyly of six major taxa was supported: Pectinariidae, Ampharetinae, Alvinellidae, Trichobranchidae, Terebellidae and Melinninae. The latter, traditionally a subfamily of Ampharetidae, was unexpectedly the sister to Terebellidae, and hence becomes Melinnidae, and Ampharetinae becomes Ampharetidae. We found no support for the recently proposed separation of Telothelepodidae, Polycirridae and Thelepodidae from Terebellidae. Telothelepodidae was nested within Thelepodinae and is accordingly made its junior synonym. Terebellidae contained the subfamily-ranked taxa Terebellinae and Thelepodinae. The placement of the simplified Polycirridae within Terebellinae differed from previous hypotheses, warranting the division of Terebellinae into Lanicini, Procleini, Terebellini and Polycirrini. Ampharetidae (excluding Melinnidae) were well-supported as the sister group to Alvinellidae and we recognize three clades: Ampharetinae, Amaginae and Amphicteinae. Our analysis found several paraphyletic genera and undescribed species. Morphological transformations on the phylogeny supported the hypothesis of an ancestor that possessed both branchiae and chaetae, which is at odds with proposals of a “naked” ancestor. Our study demonstrates how a robust backbone phylogeny can be combined with dense taxon coverage and morphological traits to give insights into the evolutionary history and transformation of traits.

Polygordius is a clade of marine annelids mainly seen in coarse-grained habitats. They are notable for their smooth bodies, lacking in chaetae or obvious segments, and they resemble Nematoda or Nemertea. Though Polygordius taxa are found in all oceans of the world, identifying species based only on morphological characters can be challenging due to their relatively uniform external appearances. Diversity within the clade has likely been markedly underestimated. Where morphological characters are inconspicuous or even unreliable, molecular methods can provide clarity in delimiting species. In this study, morphological methods (examination under light and scanning electron microscopy) were integrated with molecular analyses (sequencing of Cytochrome c oxidase subunit I, 16S rRNA and Histone H3 gene fragments) to establish the systematic placement of Polygordius specimens collected from Australia, Belize, French Polynesia, Indonesia, Japan, and the U.S. west coast. These analyses revealed three new species of Polygordius from the Pacific Ocean (P. erikae n. sp., P. kurthcarolae n. sp., and P. kurthsusanae n. sp.) as well as one new species from the Caribbean Sea (P. jenniferae n. sp.). These new species are formally described, and a previously known Japanese species, P. pacificus Uchida, 1936, is redescribed. This study establishes the first molecular data set for Polygordius species from the Pacific region, as well as the first formal description of a Caribbean species of Polygordius. Phylogenetic relationships within Polygordius are summarized and discussed.

We describe two new species of the annelid genus Trilobodrilus Remane, 1925 (Dinophilidae Verill, 1892) from an intertidal and a subtidal location in San Diego, California. These two species show morphological and molecular divergences between each other and the previously described, geographically distant species. Intertidal T. windansea sp. nov. differs from subtidal T. ellenscrippsae sp. nov. most remarkably in the number and pattern of ciliary tufts and bands on the prostomium and along the body length, besides showing ca 15% difference in gene fragments of COI and CytB. Trilobodrilus windansea sp. nov., though nesting with T. ellenscrippsae sp. nov. in the molecular phylogenetic analyses, morphologically resembles the Japanese T. itoi Kajihara, Ikoma, Yamasaki & Hiruta, 2015 most closely, but still differs from this species in the higher number of apical ciliary tufts, an additional ciliary row posterior to the second ciliary band, and by lacking a forth ciliary band and segmentally arranged lateral ciliary tufts. Trilobodrilus ellenscrippsae sp. nov. is morphologically most similar to the Japanese T. nipponicus Uchida & Okuda, 1943, but is much shorter, has more apical ciliary tufts, and less regularly arranged lateral ciliary tufts along the body. All species differ significantly in all compared gene fragments, and no obvious correlation was found between habitat and the species morphology or relationships.

The four named species of Branchipolynoe all live symbiotically in mytilid mussels (Bathymodiolus) that occur at hydrothermal vents or methane seeps. Analyses using mitochondrial (COI and 16S) and nuclear (ITS) genes, as well as morphology, were conducted on a collection of Branchipolynoe from Pacific Costa Rican methane seeps and West Pacific hydrothermal vents. This revealed five new species of Branchipolynoe, and these are formally described. The new species from Costa Rica live in three species of Bathymodiolus mussels (also new) at depths ranging from 1000 to 1800 m. Branchipolynoe kajsae n. sp. and Branchipolynoe halliseyae n. sp. were found in all three undescribed Bathymodiolus species, while Branchipolynoe eliseae n. sp. was found in Bathymodiolus spp. 1 and 2, and Branchipolynoe meridae n. sp. was found in Bathymodiolus spp. 1 and 3. Hence, Bathymodiolus sp. 1 hosted all four of the new species, while the other two Bathymodiolus hosted three each. Most mussels contained only one specimen of Branchipolynoe; where there was more than one, these were often a female and smaller male of the same species. The newly discovered species from the West Pacific, Branchipolynoe tjiasmantoi n. sp., lives in unidentified Bathymodiolus at depths ranging from 674 to 2657 m from hydrothermal vents in the North Fiji (Fiji) and Lau Basins (Tonga) and also from New Zealand, Vanuatu, and the Manus Basin (Papua New Guinea). The phylogenetic and biogeographical implications of this diversity of Branchipolynoe are discussed.

Assessing the enormous diversity of Southern Ocean benthic species and their evolutionary histories is a central task in the era of global climate change. Based on mitochondrial markers, it was recently suggested that the circumpolar giant sea spider Colossendeis megalonyx comprises a complex of at least six cryptic species with mostly small and non-overlapping distribution ranges. Here, we expand the sampling to include over 500 mitochondrial COI sequences of specimens from around the Antarctic. Using multiple species delimitation approaches, the number of distinct mitochondrial OTUs increased from six to 15–20 with our larger dataset. In contrast to earlier studies, many of these clades show almost circumpolar distributions. Additionally, analysis of the nuclear internal transcribed spacer region for a subset of these specimens showed incongruence between nuclear and mitochondrial results. These mito-nuclear discordances suggest that several of the divergent mitochondrial lineages can hybridize and should not be interpreted as cryptic species. Our results suggest survival of C. megalonyx during Pleistocene glaciations in multiple refugia, some of them probably located on the Antarctic shelf, and emphasize the importance of multi-gene datasets to detect the presence of cryptic species, rather than their inference based on mitochondrial data alone.

The exploration of Earth's biodiversity is an exciting and ongoing endeavour. Here, we report a new species of seadragon from Western Australia with substantial morphological and genetic differences to the only two other known species. We describe it as Phyllopteryx dewysea n. sp. Although the leafy seadragon (Phycodurus eques) and the common seadragon (Phyllopteryx taeniolatus) occur along Australia's southern coast, generally among relatively shallow macroalgal reefs, the new species was found more offshore in slightly deeper waters. The holotype was trawled east of the remote Recherche Archipelago in 51 m; additional specimens extend the distribution west to Perth in 72 m. Molecular sequence data show clear divergence from the other seadragons (7.4–13.1% uncorrected divergence in mitochondrial DNA) and support a placement as the sister-species to the common seadragon. Radiographs and micro-computed tomography were used on the holotype of the new species and revealed unique features, in addition to its unusual red coloration. The discovery provides a spectacular example of the surprises still hidden in our oceans, even in relatively shallow waters.