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Modern poriferans are classified into four classes-Calcarea, Demospongiae, Hexactinellida and Homoscleromorpha-the recognition of which in fossil specimens almost exclusively relies on spicule morphology and arrangement. Early fossil representatives of the phylum Porifera are morphologically diverse, and many of them problematically display characteristics that are incompatible with the classification scheme developed for modern taxa. Critically, hexactine spicules-a diagnostic feature of hexactinellids among modern taxa-are found in various Cambrian and Ordovician taxa that cannot be accommodated within the hexactinellid body plan. Here we describe a new poriferan from the Drumian Marjum Formation of Utah, Polygoniella turrelli gen. et sp. nov., which exhibits a unique combination of complex anatomical features for a Cambrian form, including a syconoid-like organization, a thick body wall, and a multi-layered hexactin-based skeleton. The hexactinellid-like body wall architecture of this new species supports a Cambrian origin of the hexactinellid body plan and provides valuable insights into character evolution in early glass sponges.

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Batellopsis paula gen. et sp. nov., a new hexactinellid-associated alpheid shrimp, is described based on a single specimen collected at a depth of 477-503 m north of Île des Pins, New Caledonia. Batellopsis gen. nov. is part of a clade of four genera all containing hexactinellid-associated species, for which a molecular phylogeny is presented. The evolution of several morphological characters, including orbital teeth, a bulgefossa system on the fingers of the first pereiopod chela, and groups of microserrulate setae on the second pereiopod chela, is discussed in light of phylogenetic results.

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Bast fibre-reinforced plastics are characterised by good strength and stiffness but are often brittle due to the stiff and less ductile fibres. This study uses a biomimetic approach to improve impact strength. Based on the structure of the spicules of a deep-sea glass sponge, in which hard layers of bioglass alternate with soft layers of proteins, the toughness of kenaf/epoxy composites was significantly improved by a multilayer structure of kenaf and cellulose acetate (CA) foils as impact modifiers. Due to the alternating structure, cracks are deflected, and toughness is improved. One to five CA foils were stacked with kenaf layers and processed to composite plates with bio-based epoxy resin by compression moulding. Results have shown a significant improvement in toughness using CA foils due to increased crack propagation. The unnotched Charpy impact strength increased from 9.0 kJ/m2 of the pure kenaf/epoxy composite to 36.3 kJ/m2 for the sample containing five CA foils. The tensile and flexural strength ranged from 74 to 81 MPa and 112 to 125 MPa, respectively. The tensile modulus reached values between 9100 and 10,600 MPa, and the flexural modulus ranged between 7200 and 8100 MPa. The results demonstrate the successful implementation of an abstract transfer of biological role models to improve the toughness of brittle bast fibre-reinforced plastics.

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Well-annotated and contiguous genomes are an indispensable resource for understanding the evolution, development, and metabolic capacities of organisms. Sponges, an ecologically important non-bilaterian group of primarily filter-feeding sessile aquatic organisms, are underrepresented with respect to available genomic resources. Here we provide a high-quality and well-annotated genome of Aphrocallistes vastus, a glass sponge (Porifera: Hexactinellida) that forms large reef structures off the coast of British Columbia (Canada). We show that its genome is approximately 80 Mb, small compared to most other metazoans, and contains nearly 2500 nested genes, more than other genomes. Hexactinellida is characterized by a unique skeletal architecture made of amorphous silicon dioxide (SiO2), and we identified 419 differentially expressed genes between the osculum, i.e. the vertical growth zone of the sponge, and the main body. Among the upregulated ones, mineralization-related genes such as glassin, as well as collagens and actins, dominate the expression profile during growth. Silicateins, suggested being involved in silica mineralization, especially in demosponges, were not found at all in the A. vastus genome and suggests that the underlying mechanisms of SiO2 deposition in the Silicea sensu stricto (Hexactinellida + Demospongiae) may not be homologous.

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Sponge fossils from the Cambrian black shales have attracted attention from both palaeontologists and geochemists for many years in terms of their high diversity, beautiful preservation and perplexing adaptation to inhospitable living environments. However, the body shape of these sponges, which contributes to deciphering adaptive evolution, has not been scrutinized. New complete specimens of the hexactinellid sponge Sanshapentella tentoriformis sp. nov. from the Qingjiang biota (black shale of the Cambrian Stage 3 Shuijingtuo Formation, ca 518 Ma) allow recognition of a unique dendriform body characterized by a columnar trunk with multiple conical high peaks and distinctive quadripod-shaped dermal spicules that frame each high peak. The body shape of this new sponge along with other early Cambrian hexactinellids, is classified into three morpho-groups that reflect different levels of adaptivity to the environment. The cylindrical and ovoid bodies generally adapted to a large spectrum of environments; however, the dendriform body of S. tentoriformis was restricted to the relatively deep-water, oxygen-deficient environment. From a hindsight view, the unique body shape represents a consequence of adaptation that helps maintain an effective use of oxygen and a low energy cost in hypoxic conditions.

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Few studies have explored the microbiomes of glass sponges (Hexactinellida). The present study seeks to elucidate the composition of the microbiota associated with the glass sponge Vazella pourtalesii and the functional strategies of the main symbionts. We combined microscopic approaches with metagenome-guided microbial genome reconstruction and amplicon community profiling toward this goal. Microscopic imaging revealed that the host and microbial cells appeared within dense biomass patches that are presumably syncytial tissue aggregates. Based on abundances in amplicon libraries and metagenomic data, SAR324 bacteria, Crenarchaeota, Patescibacteria, and Nanoarchaeota were identified as abundant members of the V. pourtalesii microbiome; thus, their genomic potentials were analyzed in detail. A general pattern emerged in that the V. pourtalesii symbionts had very small genome sizes, in the range of 0.5 to 2.2 Mb, and low GC contents, even below those of seawater relatives. Based on functional analyses of metagenome-assembled genomes (MAGs), we propose two major microbial strategies: the "givers," namely, Crenarchaeota and SAR324, heterotrophs and facultative anaerobes, produce and partly secrete all required amino acids and vitamins. The "takers," Nanoarchaeota and Patescibacteria, are anaerobes with reduced genomes that tap into the microbial community for resources, e.g., lipids and DNA, likely using pilus-like structures. We posit that the existence of microbial cells in sponge syncytia together with the low-oxygen conditions in the seawater environment are factors that shape the unique compositional and functional properties of the microbial community associated with V. pourtalesii IMPORTANCE We investigated the microbial community of V. pourtalesii that forms globally unique, monospecific sponge grounds under low-oxygen conditions on the Scotian Shelf, where it plays a key role in its vulnerable ecosystem. The microbial community was found to be concentrated within biomass patches and is dominated by small cells (<1 µm). MAG analyses showed consistently small genome sizes and low GC contents, which is unusual compared to known sponge symbionts. These properties, as well as the (facultatively) anaerobic metabolism and a high degree of interdependence between the dominant symbionts regarding amino acid and vitamin synthesis, are likely adaptations to the unique conditions within the syncytial tissue of their hexactinellid host and the low-oxygen environment.

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In the present study, we profiled bacterial and archaeal communities from 13 phylogenetically diverse deep-sea sponge species (Demospongiae and Hexactinellida) from the South Pacific by 16S rRNA-gene amplicon sequencing. Additionally, the associated bacteria and archaea were quantified by real-time qPCR. Our results show that bacterial communities from the deep-sea sponges are mostly host-species specific similar to what has been observed for shallow-water demosponges. The archaeal deep-sea sponge community structures are different from the bacterial community structures in that they are almost completely dominated by a single family, which are the ammonia-oxidizing genera within the Nitrosopumilaceae. Remarkably, the archaeal communities are mostly specific to individual sponges (rather than sponge-species), and this observation applies to both hexactinellids and demosponges. Finally, archaeal 16s gene numbers, as detected by quantitative real-time PCR, were up to three orders of magnitude higher than in shallow-water sponges, highlighting the importance of the archaea for deep-sea sponges in general.

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In this article we report on a hexactinellid sponge new to science, Rhizophyta yapensis gen. et sp. nov., which was collected from the Yap Trench in the northwestern Pacific Ocean at an abyssal depth of 4159-4779 m. Its fungus-like body form with long peduncle and absence of hypodermalia suggest placement within the euplectellid subfamily Bolosominae Tabachnick, 2002, whereas molecular phylogenetic evidence suggests that it is sister to all remaining Euplectellidae Gray, 1867. Its rhizophytous method of attachment to the substrate, hitherto unknown from bolosomine Euplectellidae, a veil of pentactins covering the peduncle, and the presence of toothed discohexasters as the only type of microscleres, clearly characterize it as a new genus. The intraspecific divergence between holotype and paratypes of the new species is examined with both morphological and molecular approaches. This report represents the first record of a hexactinellid sponge from the Yap Trench.

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Two new Hexactinellida species from the Clarion-Clipperton Zone (CCZ) in the East Pacific Ocean are described. They are the first described representatives of the genus Sympagella in this region. The new sponges were collected in 2013 during the ABYSSLINE Project´s first cruise, AB01, on board the RV Melville. The CCZ is known for its polymetallic nodules but megafaunal biodiversity is still poorly understood. Our findings suggest that the poriferan fauna of the eastern CCZ is both species rich and inadequately known, and that substantially more sampling and taxonomic studies of the CCZ sponge fauna are required to establish a megafaunal biogeography and evaluate potential extinction risks resulting from polymetallic-nodule mining.

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The new Hexactinellid sponge Chaunoplectella megapora sp. nov. reported in this study was collected from the COMRA contract area, the western part of Clarion-Clipperton Fracture Zone (CCFZ) in the eastern Pacific Ocean at a depth of 5258 m. This sponge's extraordinary multiporous body with the presence of unique codonhexasters, sigmatocomes, toothed discohexasters and hemidiscohexasters, as well as stellate disocohexasters, characterizes it as a new species in the genus Chaunoplectella. This report presents the first record of family Leucopsacidae at this site in the eastern Pacific Ocean.

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A new species of the deep-sea spongicolid genus Spongicoloides Hansen, 1908 is described and illustrated based on material from the northwestern Pacific. Spongicoloides weijiaensissp. n. was found inside a hexactinellid sponge, Euplectella sp., sampled by the Chinese manned submersible "Jiaolong" at depths of 2279 m near the Weijia Guyot, in the Magellan Seamount Chain. The new species can be distinguished from all congeneric species by several morphological features, involving gill formula, spination of the carapace, antennal scale, third pereiopod, telson and uropod, posteroventral teeth of the pleura, and dactyli of the fourth and fifth pereiopods. An identification key to the Pacific species of Spongicoloides is provided.

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BACKGROUND: Glass sponges (Class Hexactinellida) are important components of deep-sea ecosystems and are of interest from geological and materials science perspectives. The reconstruction of their phylogeny with molecular data has only recently begun and shows a better agreement with morphology-based systematics than is typical for other sponge groups, likely because of a greater number of informative morphological characters. However, inconsistencies remain that have far-reaching implications for hypotheses about the evolution of their major skeletal construction types (body plans). Furthermore, less than half of all described extant genera have been sampled for molecular systematics, and several taxa important for understanding skeletal evolution are still missing. Increased taxon sampling for molecular phylogenetics of this group is therefore urgently needed. However, due to their remote habitat and often poorly preserved museum material, sequencing all 126 currently recognized extant genera will be difficult to achieve. Utilizing morphological data to incorporate unsequenced taxa into an integrative systematics framework therefore holds great promise, but it is unclear which methodological approach best suits this task. RESULTS: Here, we increase the taxon sampling of four previously established molecular markers (18S, 28S, and 16S ribosomal DNA, as well as cytochrome oxidase subunit I) by 12 genera, for the first time including representatives of the order Aulocalycoida and the type genus of Dactylocalycidae, taxa that are key to understanding hexactinellid body plan evolution. Phylogenetic analyses suggest that Aulocalycoida is diphyletic and provide further support for the paraphyly of order Hexactinosida; hence these orders are abolished from the Linnean classification. We further assembled morphological character matrices to integrate so far unsequenced genera into phylogenetic analyses in maximum parsimony (MP), maximum likelihood (ML), Bayesian, and morphology-based binning frameworks. We find that of these four approaches, total-evidence analysis using MP gave the most plausible results concerning congruence with existing phylogenetic and taxonomic hypotheses, whereas the other methods, especially ML and binning, performed more poorly. We use our total-evidence phylogeny of all extant glass sponge genera for ancestral state reconstruction of morphological characters in MP and ML frameworks, gaining new insights into the evolution of major hexactinellid body plans and other characters such as different spicule types. CONCLUSIONS: Our study demonstrates how a comprehensive, albeit in some parts provisional, phylogeny of a larger taxon can be achieved with an integrative approach utilizing molecular and morphological data, and how this can be used as a basis for understanding phenotypic evolution. The datasets and associated trees presented here are intended as a resource and starting point for future work on glass sponge evolution.

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The Late Ordovician (Hirnantian, approximately 445 million years ago) extinction event was among the largest known, with 85% species loss [1]. Post-extinction survival faunas are invariably low diversity, especially benthic communities [2], but ecological structure was restored relatively rapidly [1]. This pattern, however, reflects organisms with robust skeletons, as only one exceptionally preserved Hirnantian fossil biota was previously known [3, 4]; in particular, almost no Hirnantian sponges have been recorded. Our study reveals an extraordinarily diverse, sponge-dominated community thriving immediately after the Hirnantian extinction in Zhejiang, South China. Several contemporaneous sites preserve a total diversity of over 75 sponge species, many with preserved soft tissues, in pronounced contrast to normal survival and early recovery faunas. This diversity is unprecedented for any Hirnantian fossil group, and the fauna provides a unique window into a post-extinction ecosystem. The sponges are often large and structurally complex and represent numerous different lineages that survived the extinction. Layers with abundant sponge remains were deposited after other mass extinctions [5, 6], suggesting a general pattern of sponge abundance during collapse of Phanerozoic marine ecosystems. It is possible that the conditions of ecological collapse increase the particulate food sources for sponges, while they themselves are relatively unaffected by the crises. Furthermore, the abundance of sponges in the Hirnantian sequence of South China may have aided post-extinction ecosystem recovery by stabilizing the sediment surface, allowing sessile suspension feeders such as brachiopods, corals, and bryozoans to recover rapidly.

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In order to identify new structures, the free fatty acids from an extract of a glass sponge Aulosaccus sp. (from the north-west Pacific) belonging to one of the least chemically investigated classes (Hexactinellida), were fractionated by RP-HPLC and analyzed by NMR spectroscopy and GC-MS of their pyrrolidine derivatives, methyl(ethyl) esters and their dimethyl disulfide adducts. One hundred and twenty-three C12-C31 acids (including nine new compounds) were detected, one hundred and ten of these compounds have not been found previously in glass sponges. The levels of common methylene-interrupted polyenes, monoenes of the (n-7) family and less common branched-chain components proved to be high. New acids were shown to be 5,13-dimethyl-tetradec-4-enoic, cis-10,11-methylene-heptadecanoic, 10,12-dimethyl-octadecanoic, cis-12,13-methylene-nonadecanoic, (14E)-13-methyl-eicos-14-enoic, 19-methyl-eicos-13-enoic, cis-20,21-methylene-heptacosanoic, 27-methyl-octacos-21-enoic and (22Z)-nonacos-22-enoic. Some important mass spectrometric characteristics of pyrrolidides of homologous cyclopropane fatty acids are reported and discussed.

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BACKGROUND: The checklist of Porifera of Greece was created in the framework of the Greek Taxon Information System (GTIS), an initiative of the LifeWatchGreece Research Infrastructure (ESFRI) that has resumed efforts to compile a complete checklist of species recorded from Greece. An updated checklist of Porifera was created on the basis of a list of the Aegean Demospongiae and Homoscleromorpha published one decade ago. All records of species known to occur in Greek waters were taxonomically validated and cross-checked for possible inaccuracies and omissions. Then, all recent publications were reviewed and the species recorded from 2006 to date were added to the list. NEW INFORMATION: The updated checklist of Porifera of Greece comprises 215 species, classified to 111 genera, 65 families, 24 orders, and 4 classes. In total, 34 new additions were made to the previous species list (8 Calcarea, 17 Demospongiae, 1 Hexactinellida, and 6 Homoscleromorpha) with Calcarea being listed for the first time from the area. The demosponge orders Poecilosclerida, Dictyoceratida, Tetractinellida, Haplosclerida, and Suberitida have the highest number of species covering 62% of the known Greek sponge species richness. It is worth mentioning that 8 species have been first described from Greek waters, 7 of which are considered endemic to this area. Our bibliographic overview also revealed knowledge gaps with regard to specific habitats typically rich in sponge diversity, and marine sectors of Greece.

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The minor cerebrosides from a Far-Eastern glass sponge Aulosaccus sp. were analyzed as constituents of some multi-component RP-HPLC fractions. The structures of eighteen new and one known cerebrosides were elucidated on the basis of NMR spectroscopy, mass spectrometry, optical rotation data and chemical transformations. These ß-D-glucopyranosyl-(1→1)-ceramides contain sphingoid bases N-acylated with straight-chain (2R)-2-hydroxy fatty acids, namely, (2S,3S,4R,11Z)-2-aminoeicos-11-ene-1,3,4-triol, acylated with 15E-22:1, 16Z-21:1, 15Z-21:1, 15Z-20:1, 15E-20:1, 19:0, 18:0 acids, (2S,3S,4R)-2-amino-13-methyltetradecane-1,3,4-triol--with 19Z-26:1, 16Z-23:1, 23:0, 22:0 acids, (2S,3S,4R)-2-amino-14-methylpentadecane-1,3,4-triol--with 16Z-23:1, 16E-23:1, 15Z-22:1, 22:0 acids, (2S,3S,4R)-2-amino-14-methylhexadecane-1,3,4-triol, linked to 16Z-23:1, 15Z-22:1 acids, (2S,3S,4R)-2-amino-9-methylhexadecane-1,3,4-triol--to 16Z-23:1 acid, and (2S,3S,4R)-2-aminohexadecane-1,3,4-triol, attached to 15Z-22:1 acid. The 13-methyl and 9-methyl-branched trihydroxy sphingoid base backbones (C15 and C17, respectively) have not been found previously in sphingolipids. The ceramide parts, containing other backbones, present new variants of N-acylation of the marine sphingoid bases with the 2-hydroxy fatty acids. The combination of the instrumental and chemical methods used in this study improved the efficiency of the structural analysis of such complex cerebroside mixtures that gave more detailed information on glycosphingolipid metabolism of the organism.

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Three previously studied mitochondrial genomes of glass sponges (phylum Porifera, class Hexactinellida) contained single nucleotide insertions in protein coding genes inferred as sites of +1 translational frameshifting. To investigate the distribution and evolution of these sites and to help elucidate the mechanism of frameshifting, we determined eight new complete or nearly complete mtDNA sequences from glass sponges and examined individual mitochondrial genes from three others. We found nine new instances of single nucleotide insertions in these sequences and analyzed them both comparatively and phylogenetically. The base insertions appear to have been gained and lost repeatedly in hexactinellid mt protein genes, suggesting no functional significance for the frameshifting sites. A high degree of sequence conservation, the presence of unusual tRNAs, and a distinct pattern of codon usage suggest the "out-of-frame pairing" model of translational frameshifting. Additionally, we provide evidence that relaxed selection pressure on glass sponge mtDNA - possibly a result of their low growth rates and deep-water lifestyle - has allowed frameshift insertions to be tolerated for hundreds of millions of years. Our study provides the first example of a phylogenetically diverse and extensive usage of translational frameshifting in animal mitochondrial coding sequences.

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New hexactinellid sponges were collected from 2589 m depth on the Carlsberg Ridge in the Indian Ocean during deep-sea dredging. All fragments belong to a new genus and species, Indiellagen. n.ridgenensissp. n., a representative of the family Aulocalycidae described here. The peculiar features of this sponge, not described earlier for other Aulocalycidae, are: longitudinal strands present in several layers and epirhyses channelization.