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Polymastiidae (Porifera: Demospongiae) of the Nordic and Siberian Seas are revised and compared with the related species of the North Atlantic based on the morphological data from the type and comparative material and the molecular data from fresh samples. Twenty species from six polymastiid genera are recorded. Two species, Polymastia svenseni from Western Norway and Spinularia njordi from the Norwegian Sea, are new to science. One species, Polymastia andrica, is new to the Nordic Seas and two species, Polymastia cf. bartletti and P. penicillus, are new to the Scandinavian Coast. Distribution of the polymastiids in the North Atlantic and Arctic is discussed and the allegedly wide distribution of Spinularia sarsii and S. spinularia is questioned. ; We would like to express our gratitude to all colleagues, who kindly provided the access to the sponge collections of their institutions: Jean-Marc Gagnon (Canadian Museum of Nature, Ottawa), Erica Mejlon (Museum of Evolution, University of Uppsala), Kennet Lundin and Carola Azurduy Högström (Gothenburg Natural History Museum), Miche`le Bruni (Musée Océanographique de Monaco), Carsten Lüter (Museum für Naturkunde, Berlin), Isabelle Domart-Coulon (Muséum National d'Histoire Naturelle, Paris), Clare Valentine, Andrew Cabrinovic and Emma Sherlock (Natural History Museum, London), Nicole de Voogd and J. Koos van Egmond (Naturalis Biodiversity center, Leiden), Lutz Bachmann and Ase Ingvild Wilhelmsen (Natural History Museum, University of Oslo), Ole Secher Tendal (Natural History Museum of Denmark, University of Copenhagen), Bernard E. Picton (Ulster Museum, National Museums of Northern Ireland, Belfast), Jon Anders Kongsrud (University Museum of Bergen, Natural History Collections), Torkild Bakken (University Museum of the Norwegian University of Science and Technology, Trondheim), Olga Bozhenova (Zoological Institute of Russian Academy of Sciences, Saint-Petersburg). We are also thankful to Bjørn Gulliksen (University of Tromsø), Bjørn Tore Dragnes (OMNIMAR Dragnes, ...

Family Cladorhizidae, comprising the carnivorous sponges, represents a unique innovation within the phylum Porifera. Rather than filter feeding, carnivorous sponges have developed the ability to passively capture small invertebrates such as crustaceans using filaments or other appendages, coupled with an adhesive surface and the ability of sponge cells to migrate to and envelop prey items. Cladorhizids are most commonly deep-sea species and are found worldwide, with around 150 species currently described. The boreal Atlantic and neighbouring Arctic areas have a species-rich cladorhizid fauna, including many of the first cladorhizids described from early marine biological investigations. While a number of records exist for parts of these areas, other areas, such as the North-west Atlantic, are less well known, and species descriptions are scattered and in many cases lacking necessary information. Using a large set of specimens from newly collected material and multiple museum collections, and integrating this with previously published records from the area, we provide an overview of the cladorhizid fauna of the boreal Atlantic and adjoining Arctic Oceans. In all, we provide updated descriptions of 25 species, of which four, Asbestopluma (Asbestopluma) ruetzleri sp. nov., Cladorhiza kenchingtonae sp. nov., Lycopodina novangliae sp. nov. and L. tendali sp. nov., are previously undescribed. We also provide an overview of the known distribution and depth ranges of each species as well as a key for identification to species level. Finally, we provide some general discussion on the boreo-Atlantic and Arctic cladorhizid fauna and its relationship to neighbouring areas. ; Numerous people have helped us for different parts of this study: We are greatly indebted to Ole S. Tendal for his help in providing access to the large number of cladorhizids found in the zoological collections at the Natural History Museum of Denmark. We would like to thank Megan Best for collecting SEM images of DFO specimens, and Ellen Kenchington ...

All polymastiid sponges displaying ornamented exotyles are reviewed and their morphological affinities are reconsidered. The study embraces all known species of Proteleia, Sphaerotylus, Trachyteleia and Tylexocladus as well as several species of Polymastia. A new genus, Koltunia, is established for the Antarctic species Proteleia burtoni based on the unique shape of distal ornamentations of its giant exotyles and on the absence of a spicule palisade in its cortex, a rare feature among the polymastiids. Three new species of Sphaerotylus are described – S. renoufi from the British Isles, S. strobilis from South Africa and S. tjalfei from West Greenland. Transfer of one New Zealand species from Polymastia to Proteleia and of one Chilean species from Polymastia to Sphaerotylus is proposed. The present study provides a background for future integrative phylogenetic analyses based on comprehensive molecular and morphological datasets which should reveal the natural relationships between the polymastiid taxa. ; We would like to express our gratitude to all colleagues who kindly provided the access to the historical and recent sponge collections of their institutions, namely to Clare Valentine, Andrew Cabrinovic and Emma Sherlock (National History Museum, London), Bruce Marshall (Museum of New Zealand Te Papa Tongarewa, Wellington), Bernard E. Picton (Ulster Museum, Belfast), Carsten Lüter and Carsten Eckert (Museum für Naturkunde, Berlin), Michèle Bruni (Musée Océanographique de Monaco), Ole Secher Tendal (Natural History Museum of Denmark, University of Copenhagen), Boris Sirenko and Olga Sheiko (Zoological Institute of Russian Academy of Sciences, Saint-Petersburg), Claude Lévi and Isabelle Domart-Coulon (Muséum National d'Histoire Naturelle, Paris), Klaus Rützler (Smithsonian National Museum of Natural History, Washington), Nicole de Voogd and J. Koos van Egmond (National Museum of Natural History, Leiden). Egil Severin Erichsen (University of Bergen) and Marie-Louise Tritz (Naturmuseum Senckenberg, Frankfurt am ...

Polymastiidae Gray, 1867 is a worldwide distributed sponge family, which has a great significance for understanding of the demosponge deep phylogeny since the former order Hadromerida Topsent, 1894 has been recently split based on the molecular evidence and a new separate order has been established for the polymastiids. However, molecular data obtained from Polymastiidae so far are scarce, while the phylogenetic reconstruction based on morphology has faced a deficit of characters along with the vagueness of their states. The present study is a phylogenetic reconstruction of Polymastiidae based on novel data on two molecular markers, cytochrome oxidase subunit I and large subunit ribosomal DNA, obtained from a broad set of species. Monophyly of the family and nonmonophyly of four polymastiid genera are revealed, suggesting a high level of homoplasy of morphological characters, which are therefore not an appropriate base for the natural classification of Polymastiidae. Although the presented phylogenies cannot yet provide an alternative classification scheme, several strongly supported clades, which may be used as reference points in future classification, are recovered and three taxonomic actions are proposed: transfer of one species from Radiella to Polymastia Bowerbank, 1862; transfer of three species from Radiella Schmidt, 1870 to Spinularia Gray, 1867; and the consequent abandonment of Radiella. ; This record in Zenodo includes three Supplementary Material files, available to authorized users also on the Journal's article page. This study was supported by the Norwegian Biodiversity Information Centre (grant to HTR, project number 70184219), the Norwegian Academy of Science and Letters (grant to HTR), the Research Council of Norway (through contract number 179560), and the SponGES project through Horizon 2020, the European Union Framework Program for Research and Innovation (grant agreement No 679849). © The Authors 2016. This article is published with open access at Springerlink.com. This article is ...

ABSTRACT. The Sognefjord is the longest (205 km) and deepest (1308 m) fjord in Norway, and the second-longest in the world. Coast-fjord exchange in Sognefjord is limited by a seaward sill at 170 m water depth, which causes a clear stratification between water masses as the dense oxygen-poor basin water mixes slowly with the well-oxygenated water directly above from the coastal ocean. Due to the homogeneity and limited variability in the deep-water, the deep slopes of Sognefjord represent the ideal setting to study how abiotic factors influence the deep-water benthic community structure. During the summer of 2017, two remotely operated vehicle (ROV) video transects were performed to compare the megabenthic community behind the sill (water depth: 1230 to 55 m; transect length: 1.39 km; distance from sill: ∼17 km) and within the central fjord (water depth: 1155–85 m; transect length: 2.43 km; distance from sill: ∼79 km). Accompanying conductivity–temperature–depth (CTD) deployments were made to measure the in situ abiotic factors and nutrient concentrations at each transect location, while the substrate characteristics (percent cover of soft and hard exposed substrate) were documented from the video footage. Here, Sognefjord's megabenthic community composition, distribution, and species richness were analyzed in relation to abiotic factors (e.g., depth, salinity, dissolved oxygen, chlorophyll a concentration, and percent cover of hard and soft substrata) within the fjord. Basin communities were homogeneous and characterized by sponges, echinoderms, and crustaceans, whereas the shallower regions were dominated by mobile scavengers. Contrary to other fjord-based studies, species richness and diversity were stable in the fjord basin and decreased with proximity to the sill, decreasing water depth, and at the boundary between intermediate and basin water. The findings demonstrate that highly stratified fjords support stable communities in their basins; however, further research is needed to investigate the influence ...

ABSTRACT. Deep-sea sponge grounds are underexplored ecosystems that provide numerous goods and services to the functioning of the deep-sea. This study assessed the prokaryotic diversity in embryos, recruits, and adults of Craniella zetlandica and Craniella infrequens, common and abundant representatives of deep-sea sponge grounds in the North Atlantic. Our results reveal that symbiont transmission in the two Craniella sponge species likely occurs vertically, as highly similar microbial consortia have been identified in adults, embryos, and recruits. Moreover, transmission electron microscopy revealed high abundances of sponge-associated microorganisms, among which Chloroflexi (SAR202) were identified as common representatives by amplicon sequencing and fluorescence in situ hybridization (FISH). Equal diversity metrices, a similar overall prokaryotic community composition and a distinct dominance of the phylum Chloroflexi within all life stages are the key findings of our analyses. Information such as presented here provide understanding on the recruitment of deep-sea sponge holobionts which is needed to develop integrated management tools of such vulnerable marine ecosystems. ; FUNDING. This study was funded by the European Union's Horizon 2020 Research and Innovation Program under Grant Agreement No. 679849 (the SponGES project). This document reflects only the authors' view and the Executive Agency for Small and Medium-sized Enterprises (EASME) is not responsible for any use that may be made of the information it contains. ACKNOWLEDGEMENTS. We express our gratitude to HR, coordinator of the H2020-SponGES project on deep-sea sponge grounds of the North Atlantic, deep-sea sponge taxonomist, explorer, mentor, and friend. We thank the crews and scientific parties of RV G.O.Sars cruises "GS2017110" and "GS2018108". On these cruises the ROV AEgir-team (University of Bergen) enabled sampling of precious samples. Ina Clefsen and Andrea Hethke provided valuable support in the lab back on land. We also thank Heikki ...

Emerald Basin on the Scotian Shelf off Nova Scotia, Canada, is home to a globally unique population of the glass sponge Vazella pourtalesi. Through the analysis of both in situ photographs and trawl catch data from annual multispecies bottom-trawl surveys, we examined community composition, species density, and abundance of epibenthos and fish associated with V. pourtalesi compared to locations without this sponge. Using generalized linear models and analysis of similarities, the importance of V. pourtalesi in enhancing species density and abundance of the associated epibenthic community was assessed against that of the hard substrate on which it settles. Our results indicated that the megafaunal assemblage associated with V. pourtalesi was significantly different in composition and higher in species density and abundance compared to locations without V. pourtalesi. Analysis of similarity of trawl catch data indicated that fish communities associated with the sponge grounds are significantly different from those without V. pourtalesi, although no species were found exclusively on the sponge grounds. Our study provides further evidence of the role played by sponge grounds in shaping community structure and biodiversity of associated deep-sea epibenthic and fish communities. The mechanism for biodiversity enhancement within the sponge grounds formed by V. pourtalesi is likely the combined effect of both the sponge itself and its attachment substrate, which together comprise the habitat of the sponge grounds. We also discuss the role of habitat provision between the mixed-species tetractinellid sponges of the Flemish Cap and the monospecific glass sponge grounds of Emerald Basin. ; ACKNOWLEDGEMENTS. This work was funded in part through the SponGES—Deep-sea Sponge Grounds Ecosystems of the North Atlantic: an integrated approach towards their preservation and sustainable exploitation, under H2020 - the EU Framework Programme for Research and Innovation (Grant Agreement no. 679849) where Fisheries and Oceans Canada ...

ABSTRACT Background: Lithistid demosponges, also known as rock sponges, are a polyphyletic group of sponges which are widely distributed. In the Northeast Atlantic (NEA), 17 species are known and the current knowledge on their distribution is mainly restricted to the Macaronesian islands. In the Mediterranean Sea, 14 species are recorded and generally found in marine caves. Methods: Lithistids were sampled in nine NEA seamounts during the scientific expeditions Seamount 1 (1987) and Seamount 2 (1993) organized by the MNHN of Paris. Collected specimens were identified through the analyses of external and internal morphological characters using light and scanning electron microscopy, and compared with material from various museum collections as well as literature records. Results: A total of 68 specimens were analysed and attributed to 17 species across two orders, seven families, and seven genera, representing new records of distribution. Ten of these species are new to science, viz. Neoschrammeniella inaequalis sp. nov., N. piserai sp. nov., N. pomponiae sp. nov., Discodermia arbor sp. nov., D. kellyae sp. nov., Macandrewia schusterae sp. nov., M. minima sp. nov., Exsuperantia levii sp. nov., Leiodermatium tuba sp. nov. and Siphonidium elongatus sp. nov., and are here described and illustrated. New bathymetric records were also found for D. ramifera, D. verrucosa and M. robusta. The Meteor seamount group has a higher species richness (15 species) compared to the Lusitanian seamount group (six species). The majority of the species had their distribution restricted to one seamount, and ten are only known from a single locality, but this can be a result of sample bias. Discussion: The number of species shared between the seamounts and the Macaronesian islands is very reduced. The same pattern repeats between the NEA and Mediterranean Sea. This study demonstrates that NEA seamounts are ecosystems with a higher diversity of lithistids than previously thought, increasing the number of lithistids known to occur in the ...

ABSTRACT All animals are capable of undergoing gametogenesis. The ability of forming haploid cells from diploid cells through meiosis and recombination appeared early in eukaryotes, whereas further gamete differentiation is mostly a metazoan signature. Morphologically, the gametogenic process presents many similarities across animal taxa, but little is known about its conservation at the molecular level. Porifera are the earliest divergent animals and therefore are an ideal phylum to understand evolution of the gametogenic toolkits. Although sponge gametogenesis is well known at the histological level, the molecular toolkits for gamete production are largely unknown. Our goal was to identify the genes and their expression levels which regulate oogenesis and spermatogenesis in five gonochoristic and oviparous species of the genus Geodia, using both RNAseq and proteomic analyses. In the early stages of both female and male gametogenesis, genes involved in germ cell fate and cell-renewal were upregulated. Then, molecular signals involved in retinoic acid pathway could trigger the meiotic processes. During later stages of oogenesis, female sponges expressed genes involved in cell growth, vitellogenesis, and extracellular matrix reassembly, which are conserved elements of oocyte maturation in Metazoa. Likewise, in spermatogenesis, genes regulating the whole meiotic cycle, chromatin compaction, and flagellum axoneme formation, that are common across Metazoa were overexpressed in the sponges. Finally, molecular signals possibly related to sperm capacitation were identified during late stages of spermatogenesis for the first time in Porifera. In conclusion, the activated molecular toolkit during gametogenesis in sponges was remarkably similar to that deployed during gametogenesis in vertebrates. ; ACKNOWLEDGEMENTS We would like to thank Dr Nathan Kenny for his useful bioinformatic advice and Dr Cristina Díez-Vives for her useful advice in R. Furthermore, we would like to thank María Conejero for her important help in ...

Sponges occur ubiquitously in the marine realm and in some deep-sea areas they dominate the benthic communities forming complex biogenic habitats â sponge grounds, aggregations, gardens and reefs. However, deep-sea sponges and spongegrounds are still poorly investigated with regards to biotechnological potential in support of a Blue growth strategy. Under the scope of this study, five dominant North Atlantic deep-sea sponges, were characterized to elucidate promising applications in human health, namely for bone tissue engineering approaches. Geodia barretti (Gb), Geodia atlantica (Ga), Stelletta normani (Sn), Phakellia ventilabrum (Pv), and Axinella infundibuliformis (Ai), were morphologically characterized to assess macro and microstructural features, as well as chemical composition of the skeletons, using optical and scanning electron microscopy, energy dispersive x-ray spectroscopy and microcomputed tomography analyses. Moreover, compress tests were conducted to determine the mechanical properties of the skeletons. Results showed that all studied sponges have porous skeletons with porosity higher than 68%, pore size superior than 149 mm and higher interconnectivity (>96%), thus providing interesting models for the development of scaffolds for tissue engineering. Besides that, EDS analyses revealed that the chemical composition of sponges, pointed that demosponge skeletons are mainly constituted by carbon, silicon, sulfur, and oxygen combined mutually with organic and inorganic elements embedded its internal architecture that can be important features for promoting bone matrix quality and bone mineralization. Finally, the morphological, mechanical, and chemical characteristics here investigated unraveled the potential of deep-sea sponges as a source of biomaterials and biomimetic models envisaging tissue engineering applications for bone regeneration. ; The authors would like to acknowledge the funding from the European Union Framework Program for Research and Innovation Horizon 2020 through project ...

ABSTRACT Few studies have described the effects of physical disturbance and post-recovery of deep-sea benthic communities. Here, we explore the status of deep-sea sponge ground communities four years after being impacted by an experimental bottom trawl. The diversity and abundance of epibenthic megafauna of two distinct benthic communities in disturbed versus control areas were surveyed using a remotely operated vehicle on the Schulz Bank, Arctic Ocean. Four years after disturbance, megafaunal densities of the shallow (∼600 m depth) and deep (∼1,400 m depth) sites were significantly lower on the disturbed patches compared to the control areas. Multivariate analyses revealed a distinct separation between disturbed and control communities for both sites, with trawling causing 29–58% of the variation. Many epibenthic morphotypes were significantly impacted by the trawl, including ascidians, Geodia parva, Hexactinellida spp., Craniella infrequens, Lissodendoryx complicata, Haliclonia sp. Stylocordyla borealis, Gersemia rubiformis and Actiniaria sp. However, we found some smaller morphospecies to be equally abundant with control transects, including Polymastia thielei, Geodia hentscheli, and Stelletta rhaphidiophora, reflecting lower trawl impact for these morphotypes. Overall, our results suggest that these are fragile ecosystems that require much more time than four years to recover from physical disturbance typical of trawling activities. ; ACKNOWLEDGEMENTS Thanks are due to the SponGES team for their guidance, advice on our study design, and help with epifaunal morphospecies identification in the image data. Thanks to the IMAR staff for providing office space and resources. And thanks to the G.O.Sars crew, AEGIR6000 crew, and science team that carried out the ROV surveys. Finally, we would like to dedicate this contribution to our friend and colleague Hans Tore Rapp. AUTHOR CONTRIBUTIONS CP and KM conceived and carried out the study. HM, ER, and HR advised on identification of species and analysis methods. KM ...

ABSTRACT. Seamounts represent ideal systems to study the influence and interdependency of environmental gradients at a single geographic location. These topographic features represent a prominent habitat for various forms of life, including microbiota and macrobiota, spanning benthic as well as pelagic organisms. While it is known that seamounts are globally abundant structures, it still remains unclear how and to which extent the complexity of the sea floor is intertwined with the local oceanographic mosaic, biogeochemistry, and microbiology of a seamount ecosystem. Along these lines, the present study aimed to explore whether and to what extent seamounts can have an imprint on the microbial community composition of seawater and of sessile benthic invertebrates, sponges. For our high-resolution sampling approach of microbial diversity (16S rRNA gene amplicon sequencing) along with measurements of inorganic nutrients and other biogeochemical parameters, we focused on the Schulz Bank seamount ecosystem, a sponge ground ecosystem which is located on the Arctic Mid-Ocean Ridge. Seawater samples were collected at two sampling depths (mid-water, MW, and near-bed water, BW) from a total of 19 sampling sites. With a clustering approach we defined microbial microhabitats within the pelagic realm at Schulz Bank, which were mapped onto the seamount's topography and related to various environmental parameters (such as suspended particulate matter, SPM; dissolved inorganic carbon, DIC; silicate, SiO−4; phosphate, PO3−4; ammonia, NH+4; nitrate, NO2−3; nitrite, NO−2; depth; and dissolved oxygen, O2). The results of our study reveal a "seamount effect" (sensu stricto) on the microbial mid-water pelagic community at least 200 m above the sea floor. Further, we observed a strong spatial heterogeneity in the pelagic microbial landscape across the seamount, with planktonic microbial communities reflecting oscillatory and circulatory water movements, as well as processes of bentho-pelagic coupling. Depth, NO2−3, SiO−4, and O2 ...

An Open Access paper submitted under the category "Uncovered". ; We acknowledge the financial support from European Union H2020 programme (SponGES – 679849, CHEM2NATURE – 692333 and FORECAST – 668983) and FP7 programme (ERC Advanced Grant COMPLEXITE, ERC- 2012-ADG-20120216-321266).

Demosponges are a well-known source of a plethora of bioactive compounds. In particular, they are able to form a skeleton by direct deposition of silica in a process catalysed by silicatein. Herein, we isolated biosilicas from five different Atlantic deep-sea sponges Geodia atlantica (GA), Geodia barretti (GB), Stelletta normani (SN), Axinella infundibuliformis (AI) and Phakellia ventilabrum (PV) to explore the bioactivity and osteogenic capacity of its silica-based materials. We chemically characterized the isolated biosilicas and evaluated them for their bioactivity to deposit Ca and P on their surface (by immersion in simulated body fluid, SBF). GB-, SN-, AI- and PV-based biosilicas did not generate a stable calcium phosphate (CaP) layer over time in the presence of SBF, however, the GA-derived one was able to form a CaP surface layer (at a Ca/P ratio of ~1.7, similar to the one observed for hydroxyapatite), that was stable during the 28 days of testing. In addition, no cytotoxicity towards L929 and SaOs2 cells was observed for the GA-based biosilica up to a concentration of 10 mg/mL. Overall, the GA-based biosilica presents the characteristics to be used in the development of biomaterials for bone tissue engineering (BTE). ; This research has been performed with funding from the European Union's Horizon 2020 research and innovation programme under Grant Agreements No. 679849 (SponGES) and No. 668983 (FORECAST). Funding from the European Regional Development Fund, through Atlantic Area Program, under the scope of BlueHuman project (EAPA_151/2016) is also acknowledged. JRX research was also supported by National Funds through FCT, the Portuguese Foundation for Science and Technology within the scope of UIDB/04423/2020, UIDP/04423/2020, and ...

ABSTRACT. Sponges are commonly known as general nutrient providers for the marine ecosystem, recycling organic matter into various forms of bioavailable nutrients such as ammonium and nitrate. In this study we challenge this view. We show that nutrient removal through microbial denitrification is a common feature in six cold-water sponge species from boreal and Arctic sponge grounds. Denitrification rates were quantified by incubating sponge tissue sections with 15NO−3 - amended oxygen-saturated seawater, mimicking conditions in pumping sponges, and de-oxygenated seawater, mimicking non-pumping sponges. It was not possible to detect any rates of anaerobic ammonium oxidation (anammox) using incubations with 15NH+4. Denitrification rates of the different sponge species ranged from below detection to 97 nmol N cm−3 sponge d−1 under oxic conditions, and from 24 to 279 nmol N cm−3 sponge d−1 under anoxic conditions. A positive relationship between the highest potential rates of denitrification (in the absence of oxygen) and the species-specific abundances of nirS and nirK genes encoding nitrite reductase, a key enzyme for denitrification, suggests that the denitrifying community in these sponge species is active and prepared for denitrification. The lack of a lag phase in the linear accumulation of the 15N-labelled N2 gas in any of our tissue incubations is another indicator for an active community of denitrifiers in the investigated sponge species. Low rates for coupled nitrification–denitrification indicate that also under oxic conditions, the nitrate used to fuel denitrification rates was derived rather from the ambient seawater than from sponge nitrification. The lack of nifH genes encoding nitrogenase, the key enzyme for nitrogen fixation, shows that the nitrogen cycle is not closed in the sponge grounds. The denitrified nitrogen, no matter its origin, is then no longer available as a nutrient for the marine ecosystem. These results suggest a high potential denitrification capacity of deep-sea sponge grounds ...