Diversity, habitat endemicity and trophic ecology of the fauna of Loki's Castle vent field on the Arctic Mid-Ocean Ridge.

Loki's Castle Vent Field (LCVF, 2300 m) was discovered in 2008 and represents the first black-smoker vent field discovered on the Arctic Mid-Ocean Ridge (AMOR). However, a comprehensive faunal inventory of the LCVF has not yet been published, hindering the inclusion of the Arctic in biogeographic analyses of vent fauna. There is an urgent need to understand the diversity, spatial distribution and ecosystem function of the biological communities along the AMOR, which will inform environmental impact assesments of future deep-sea mining activities in the region. Therefore, our aim with this paper is to provide a comprehensive inventory of the fauna at LCVF and present a first insight into the food web of the vent community. The fauna of LCVF has a high degree of novelty, with five new species previously described and another ten new species awaiting formal description. Most of the new species from LCVF are either hydrothermal vent specialists or have been reported from other chemosynthesis-based ecosystems. The highest taxon richness is found in the diffuse venting areas and may be promoted by the biogenic habitat generated by the foundation species Sclerolinum contortum. The isotopic signatures of the vent community of LCVF show a clear influence of chemosynthetic primary production on the foodweb. Considering the novel and specialised fauna documented in this paper, hydrothermal vents on the AMOR should be regarded as vulnerable marine ecosystems and protective measures must therefore be implemented, especially considering the potential threat from resource exploration and exploitation activities in the near future.

The Nereididae (Annelida) - diagnoses, descriptions, and a key to the genera.

Nereididae is among the most familiar of marine annelid families, common and well-studied in most marine environments but paradoxically no recent key or identification guide exists to the world's genera. Here updated generic descriptions, a list of characters, a linear key to genera, and minimal diagnoses that distinguish each genus from all others in the family are provided. This information is generated from a Delta database of 186 morphological characters and a link is provided to downloadable software allowing the same data to be interrogated using the open-source Delta program Intkey - a nonlinear multiple entry point computerised interactive key. For each genus the recent literature is also summarised, comments on taxonomic status provided, and published keys to species cited. Nexus format matrices are provided for all 45 genera and 158 Nereididae species, representing all genera, scored for 146 multistate characters from the same character list to facilitate future phylogenetic studies.

A regionally coherent ecological fingerprint of climate change, evidenced from natural history collections.

Climate change has dramatic impacts on ecological systems, affecting a range of ecological factors including phenology, species abundance, diversity, and distribution. The breadth of climate change impacts on ecological systems leads to the occurrence of fingerprints of climate change. However, climate fingerprints are usually identified across broad geographical scales and are potentially influenced by publication biases. In this study, we used natural history collections spanning over 250 years, to quantify a range of ecological responses to climate change, including phenology, abundance, diversity, and distributions, across a range of taxa, including vertebrates, invertebrates, plants, and fungi, within a single region, Central Norway. We tested the hypotheses that ecological responses to climate change are apparent and coherent at a regional scale, that longer time series show stronger trends over time and in relation to temperature, and that ecological responses change in trajectory at the same time as shifts in temperature. We identified a clear regional coherence in climate signal, with decreasing abundances of limnic zooplankton (on average by 7691 individuals m-3 °C-1) and boreal forest breeding birds (on average by 1.94 territories km-2 °C-1), and earlier plant flowering phenology (on average 2 days °C-1) for every degree of temperature increase. In contrast, regional-scale species distributions and species diversity were largely stable. Surprisingly, the effect size of ecological response did not increase with study duration, and shifts in responses did not occur at the same time as shifts in temperature. This may be as the long-term studies include both periods of warming and temperature stability, and that ecological responses lag behind warming. Our findings demonstrate a regional climate fingerprint across a long timescale. We contend that natural history collections provide a unique window on a broad spectrum of ecological responses at timescales beyond most ecological monitoring programs. Natural history collections are thus an essential source for long-term ecological research.

Describing the hidden species diversity of Chaetozone (Annelida, Cirratulidae) in the Norwegian Sea using morphological and molecular diagnostics.

Using molecular markers and species delimitation analyses, a high diversity of bi-tentaculate Cirratulidae was discovered from the North-East Atlantic. Five new species are described: Chaetozone pseudosetosa sp. nov., Chaetozone quinta sp. nov., Chaetozone barentsensis sp. nov., Chaetozone monteverdii sp. nov., and Chaetozone chambersae sp. nov. Several morphogroups are also described, even though the presence of cryptic diversity prevented naming of individual species. For each species presented, a molecular diagnostic is given from the universal barcode COI and, when available, the D1-D2 domains of the 28S rRNA. This increases the number of species in Chaetozone in northern European waters from ten to at least 17 species, the exact number of species remaining uncertain as taxonomic issues with older names remain unresolved.

Species delimitation analyses of NE Atlantic Chaetozone (Annelida, Cirratulidae) reveals hidden diversity among a common and abundant marine annelid.

The polychaetes of the family Cirratulidae (Annelida) are common inhabitants in continental shelf benthic environments and considered an important group of organisms in environmental monitoring surveys. The family represents a taxonomic and systematic challenge, as monophyly of genera and evolutionary relationships within the family remain to be explored in a proper phylogenetic framework. Bitentaculate cirratulids, especially the genus Chaetozone, form one of the most species-diverse group of polychaetes worldwide. In this study, we aimed at evaluating the species diversity of the genus Chaetozonein benthic environments in the North East Atlantic by molecular means. We tested whether traditional morphological diagnostic characters are able to discriminate between the species hypothesis after species delimitation analyses, and assessed monophyly of the genera involved. Two DNA markers were sequenced from about 200 specimens belonging to Chaetozone, Aphelochaeta, Dodecaceria, Cirriformia and Cirratulus - the universal mitochondrial barcoding region COI, and the D1-D2 regions of the nuclear 28S rRNA - and analyzed with Bayesian inference, Maximum Likelihood and the species delimitation methods mPTP and GMYC. The first phylogeny of the family Cirratulidae is inferred and the genera Chaetozone, Dodecaceria and Cirratulus are recovered monophyletic. A total of 14 clusters of sequences - corresponding to species of Chaetozone - were found in the study area, and only one of them is here referred to a nominal species, Chaetozone setosa. Our results reveal several species complexes in the genus Chaetozone, that some of these independent lineages are unnamed and undescribed, and that morphological diagnostic features are in most cases unable to discriminate between the most similar species.

Knock knock, who's there?: marine invertebrates in tubes of Ceriantharia (Cnidaria: Anthozoa).

This study reports on the fauna found in/on tubes of 10 species of Ceriantharia and discusses the characteristics of these occurrences, as well as the use of mollusc shells in ceriantharian tube construction. A total of 22 tubes of Ceriantharia from Argentina, Brazil, Japan, Norway, Portugal and the United States were analysed, revealing 58 species of marine invertebrates using them as alternative substrates. Based on a literature review and analyses of the sampled material, we report new occurrences for Photis sarae (Crustacea), Microgaza rotella (Mollusca), Brada sp., Dipolydora spp., Notocirrus spp., and Syllis garciai (Annelida). The use of mollusc shells in tube construction increases the tubes' structural resistance and strength. Ceriantharian tubes are suitable alternative substrates for the dwelling of numerous tubicolous and infaunal species that usually burrow into sediments or anchor on fixed or mobile habitats seeking shelter, thus playing a relevant role as local biodiversity hotspots.

Systematic re-structure and new species of Sphaerodoridae (Annelida) after morphological revision and molecular phylogenetic analyses of the North East Atlantic fauna.

Detailed morphological study of more than 2600 North East Atlantic (NEA) sphaerodorids (SphaerodoridaeAnnelida) and phylogenetic analyses of DNA sequences of representatives of several identified morphospecies enforced changing the current systematic classification within the family allowed the discovery of new species provided new information about the morphological and genetic characterisation of members of this group and increased the species occurrence data to better infer their geographic and bathymetric distribution ranges. Phylogenetic analyses of nuclear (18S rRNA and 28S rRNA) and mitochondrial sequences (COI and 16S rRNA) of NEA short-bodied sphaerodorids revealed outstanding results including paraphyly of the genera Sphaerodoropsis Sphaerodoridium, and Sphaerephesia. The number of longitudinal and transverse rows of dorsal macrotubercles is proposed as potential synapomorphies for the main clades and are consequently herein used for the genera delimitation. The new classification proposed here implies nomenclatural changes and the erection of a new genus Geminofilum gen. n. to accommodate the species previously considered as Sphaerodoropsis with two transverse rows of dorsal macrotubercles per segment. Four species are being described herein: Euritmianordica Capa & Bakken sp. n. Sphaerephesiamultichaeta Capa Moreira & Parapar sp. n. Sphaerephesiaponsi Capa Parapar & Moreira sp. n. and Sphaerodoridiumceliae Moreira Capa & Parapar sp. n. Characterisation of the other 21 species including updated iconography and an identification key to all NEA short-bodied sphaerodorids are provided.

Three, two, one! Revision of the long-bodied sphaerodorids (Sphaerodoridae, Annelida) and synonymization of Ephesiella, Ephesiopsis and Sphaerodorum.

BACKGROUND: Long-bodied sphaerodorids (Annelida, Sphaerodoridae) is the common name for members of the three closely and morphologically homogenous currently accepted genera of benthic marine bristle worms: Ephesiella, Ephesiopsis and Sphaerodorum. Members of this group share the presence of two dorsal and longitudinal rows of macrotubercles with terminal papillae, and two longitudinal rows of microtubercles, features that are unique among sphaerodorids. Genera are distinguished by the chaetae morphology. Members of Ephesiella are characterised by having compound chaetae (except, sometimes, simple chaetae in the first chaetigers), Sphaerodorum bear only simple chaetae, and Ephesiopsis have both compound and simple chaetae in all parapodia. METHODS: Mitochondrial (partial COI and 16S rDNA) and nuclear (partial 18S rDNA and 28S rDNA) sequence data of long-bodied sphaerodorids with compound and simple chaetae, and an outgroup of additional seven sphaerodorid species were analysed separately and in combination using Bayesian inference (BA), and Maximum Likelihood (ML) methods. Long-bodied sphaerodorids from around the world (including type specimens) were examined under a range of optical equipment in order to evaluate putative generic and specific diagnostic features, in addition to intraspecific variability. RESULTS: Phylogenetic analyses of mitochondrial and nuclear DNA sequences of specimens identified as Ephesiella and Sphaerodorum, based on chaeta morphology, were performed. Sphaerodorum and Ephesiella were recovered as paraphyletic and nested within each other. Revision of current nominal species diagnostic features are performed and discussed. DISCUSSION: Results contradict current generic definitions. Recovery of paraphyletic compound and simple chaetae clades urge the synonymization of these two genera of long-bodied sphaerodorids. Morphological data also suggest the synonymization of Ephesiopsis.

Revision of the Alitta virens species complex (Annelida: Nereididae) from the North Pacific Ocean.

Alitta virens species complex encompasses elongate nereidids appreciated commercially both in the fishing and aquaculture industries. This complex has been well studied in biological and ecological terms. Nevertheless, detailed taxonomic analyses have scarcely been addressed, to the extent that only a few species in the complex have been recognized as valid but with some difficulties: Alitta brandti Malmgren, 1865 (Sea of Okhotsk), A. grandis (Stimpson, 1853) (northeastern USA) and A. virens (Sars, 1835) (Norway). Whereas, other several species have typically been regarded as synonyms, including those originally described from the North Pacific Ocean: Nereis (Alitta) virens plenidentata Moore, 1909 (California, USA), Nereis dyamusi Izuka, 1912 (Japan) and Nereis foliata Baird, 1863 (Vancouver, Canada). In this study, an examination of the immature and epitoke type and non-type material available for the A. virens species complex from the North Pacific was carried out. Herein, the status of A. brandti as a valid species is reinforced being clearly distinct from A. virens and related species. Alitta dyamusi n. comb., A. plenidentata n. comb. and A. williami nom. nov. are reinstated and transferred to Alitta, the specific epithet of the latter species is a replacement name for Nereis foliata Baird, 1863, which is a junior, primary homonym of Nereis foliata Dalyell, 1853. Lectotypes for A. plenidentata n. comb. and A. williami nom. nov. are designated. The North Pacific species of the A. virens complex, excluding A. plenidentata n. comb. which has several distinctive features that differ from all the Alitta species, are characterized by having homogomph spinigers in both supracicular and subacicular neurochaetae, oral ring with a larger number of rows and paragnaths, and epitoke males with unmetamorphosed pygidium and epitoke-modified chaetae in both neuropodial fascicles. The morphology of epitoke females in the A. virens complex is described for the first time. Identification keys to atoke and epitoke species of this complex are also provided.

Discovery of a silicate rock-boring organism and macrobioerosion in fresh water.

Macrobioerosion is a common process in marine ecosystems. Many types of rock-boring organisms break down hard substrates, particularly carbonate rocks and calcareous structures such as dead corals and shells. In paleontology, the presence of rocks with boreholes and fossil macroboring assemblage members is one of the primary diagnostic features of shallow marine paleo-environments. Here we describe a silicate rock-boring organism and an associated community in submerged siltstone rock outcrops in Kaladan River, Myanmar. The rock-boring mussel Lignopholas fluminalis is a close relative of the marine piddocks, and its borings belong to the ichnospecies Gastrochaenolites anauchen. The neotectonic uplift of the area leading to gradual decrease of the sea level with subsequent shift from estuarine to freshwater environment was the most likely driver for the origin of this community. Our findings highlight that rocks with macroborings are not an exclusive indicator of marine paleo-ecosystems, but may also reflect freshwater habitats.

A mega-cryptic species complex hidden among one of the most common annelids in the North East Atlantic.

We investigate mitochondrial (COI, 16S rDNA) and nuclear (ITS2, 28S rDNA) genetic structure of North East Atlantic lineages of Terebellides, a genus of sedentary annelids mainly inhabiting continental shelf and slope sediments. We demonstrate the presence of more than 25 species of which only seven are formally described. Species boundaries are determined with molecular data using a broad range of analytical methods. Many of the new species are common and wide spread, and the majority of the species are found in sympatry with several other species in the complex. Being one of the most regularly encountered annelid taxa in the North East Atlantic, it is more likely to find an undescribed species of Terebellides than a described one.

Polyphyly of the traditional family Flabellinidae affects a major group of Nudibranchia: aeolidacean taxonomic reassessment with descriptions of several new families, genera, and species (Mollusca, Gastropoda).

The Flabellinidae, a heterogeneous assembly of supposedly plesiomorphic to very derived sea slug groups, have not yet been addressed by integrative studies. Here novel material of rarely seen Arctic taxa as well as North Atlantic, North and South Pacific, and tropical Indo-West Pacific flabellinid species is investigated morpho-anatomically and with multi-locus markers (partial COI, 16S rDNA, 28S rDNA and H3) which were generated and analysed in a comprehensive aeolid taxon sampling. It was found that the current family Flabellinidae is polyphyletic and its phylogeny and taxonomic patterns cannot be understood without considering members from all the Aeolidacean families and, based on a robust phylogenetic hypothesis, morpho-anatomical evolution of aeolids is more complex than suspected in earlier works and requires reclassification of the taxon. Morphological diversity of Flabellinidae is corroborated by molecular divergence rates and supports establishing three new families (Apataidae fam. n., Flabellinopsidae fam. n., Samlidae fam. n.), 16 new genera, 13 new species, and two new subspecies among the former Flabellinidae. Two families, namely Coryphellidae and Paracoryphellidae, are restored and traditional Flabellinidae is considerably restricted. The distinctness of the recently described family Unidentiidae is confirmed by both morphological and molecular data. Several species complexes among all ex-"Flabellinidae" lineages are recognised using both morphological and molecular data. The present study shows that Facelinidae and Aeolidiidae, together with traditional "Tergipedidae", deeply divide traditional "Flabellinidae." Diagnoses for all aeolidacean families are therefore provided and additionally two new non-flabellinid families (Abronicidae fam. n. and Murmaniidae fam. n.) within traditional tergipedids are established to accommodate molecular and morphological disparity. To address relationships and disparity, we propose a new family system for aeolids. Here the aeolidacean species are classified into at least 102 genera and 24 families. Operational rules for integration of morphological and molecular data for taxonomy are suggested.

Corrigenda: Polyphyly of the traditional family Flabellinidae affects a major group of Nudibranchia: aeolidacean taxonomic reassessment with descriptions of several new families, genera, and species (Mollusca, Gastropoda). https://doi.org/10.3897/zookeys.717.21885.

[This corrects the article DOI: 10.3897/zookeys.717.21885.].

Sphaerodoridae (Annelida) of the deep Northwestern Atlantic, including remarkable new species of Euritmia and Sphaerephesia.

Sphaerodoridae (Annelida) is a seeming uncommon and minimally diverse group of polychaetes in the northwestern Atlantic, with only seven species reported from the United States, and none from the eastern coast of Canada, before the present study. Review of the large Smithsonian collection (National Museum of Natural History, Washington) revealed the presence of two morphologically extraordinary undescribed species and added a new record to the north-western Atlantic region. Euritmia carolensis sp. n. is characterised by bearing approximately 20 sessile spherical papillae arranged in three transverse rows per segment, ventrum with 4-6 larger papillae near the parapodial bases and parapodia without papillae; bearing 4-5 simple chaetae that are enlarged subdistally. Sphaerephesia amphorata sp. n. is distinguished from other congeners in the presence of four longitudinal rows of sessile, bottle-shaped macrotubercles with exceptionally long digitiform terminal papilla, and parapodia with four rounded and small papillae, bearing 4-7 compound chaetae, with blades 7-11 times as long as wide. Other encountered species are also herein re-described, including intraspecific variation and updated iconography. Comparison of material also allowed some systematic changes in the group, including the synonymisation of the genus Amacrodorum with Euritmia, and the transfer of Ephesiopsis shivae to Ephesiella. A key to the species reported from the Northwestern Atlantic is provided.

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.

Revision of the Australian Sphaerodoridae (Annelida) including the description of four new species.

A revision of the complete sphaerodorid (Sphaerodoridae, Annelida) collections housed in the three major Australian museums (The Australian Museum, Museum and Art Gallery of the Northern Territory and Museum Victoria) has been performed. Specimens of three of the four species described to date from Australian waters, Ephesiella australiensis, Sphaerodoropsis exmouthensis and Sphaerodoropsis fauchaldi, have been re-encountered, resulting in changes to their previously reported distribution range. Four additional species are herein described as new: Sphaerephesia hutchingsae n. sp., Sphaerodoropsis longofalcigera n. sp., Sphaerodoropsis megatuberculata n. sp. and Sphaerodoropsis wilsoni n. sp. Moreover, Sphaerodoropsis multipapillata heteropapillata is elevated to the rank of species. A formal description of other specimens, most likely belonging to undescribed species, is not possible due to insufficient material, but information about some of their morphological features is provided. Descriptions, iconography, geographical and ecological information of all species part of this revision is provided together with a key for identification to all Australian species. An evaluation of some of the traditional generic taxonomic features is made, considering the variation observed within the Australian species.

Occurrence and distribution of Pseudoscalibregma and Scalibregma (Annelida, Scalibregmatidae) in the deep Nordic Seas, with the description of Scalibregma hanseni n. sp.

Until recent years, only a few scalibregmatid species have been known from the Nordic Seas, largely from shelf and coastal waters. Access to a large collection from deep areas has made it possible to provide more knowledge on the diversity of this group in the area. Pseudoscalibregma parvum (Hansen, 1879) is here redescribed. The species has a wide geographic distribution in the Nordic Seas, the Barents Sea, and the Kara Sea. Type specimens of Eumenia longisetosa Théel, 1879 were found to be similar to specimens of P. parvum, confirming the synonymy of the species. A new species, Scalibregma hanseni n. sp., is described from specimens found on the continental slope. It is particularly characterised by having three pairs of rather simple branchiae. Both P. parvum and S. hanseni have small spines in the most anterior chaetiger(s), resembling spines reported from a few other Pseudoscalibregma and Scalibregma species and supporting the need to emend the genus diagnosis of Pseudoscalibregma. Scalibregma abyssorum Hansen, 1879 was reassessed and considered to be a nomen dubium. Scalibregma inflatum, which has a wide distribution along the Norwegian coast and continental shelf, is found to be restricted to depths above about 900 m. Depths from 600-800 m on the continental slope represent a transition zone with fluctuations between temperate North Atlantic water (about 7°C) and cold Norwegian Sea water (below 0°C). The three species coexist in this zone, whereas P. parvum and S. hanseni n. sp. extend down to 1700 and 1200 m, respectively, on the slope at temperatures below 0°C.