The systematics of carnivorous sponges.

Carnivorous sponges are characterized by their unique method of capturing mesoplanktonic prey coupled with the complete or partial reduction of the aquiferous system characteristic of the phylum Porifera. Current systematics place the vast majority of carnivorous sponges within Cladorhizidae, with certain species assigned to Guitarridae and Esperiopsidae. Morphological characters have not been able to show whether this classification is evolutionary accurate, and whether carnivory has evolved once or in several lineages. In the present paper we present the first comprehensive molecular phylogeny of the carnivorous sponges, interpret these results in conjunction with morphological characters, and propose a revised classification of the group. Molecular phylogenies were inferred using 18S rDNA and a combined dataset of partial 28S rDNA, COI and ALG11 sequences. The results recovered carnivorous sponges as a clade closely related to the families Mycalidae and Guitarridae, showing family Cladorhizidae to be monophyletic and also including carnivorous species currently placed in other families. The genus Lycopodina is resurrected for species currently placed in the paraphyletic subgenus Asbestopluma (Asbestopluma) featuring forceps spicules and lacking sigmas or sigmancistras. The genera Chondrocladia and Cladorhiza are found to be monophyletic. However, results indicate that the subgenus Chondrocladia is polyphyletic with respect to the subgenera Meliiderma and Symmetrocladia. Euchelipluma, formerly Guitarridae, is retained, but transferred to Cladorhizidae. The four known carnivorous species currently in Esperiopsis are transferred to Abyssocladia. Neocladia is a junior homonym and is here renamed Koltunicladia. Our results provide strong evidence in support of the hypothesis that carnivory in sponges has evolved only once. While spicule characters mostly reflect monophyletic groups at the generic level, differences between genera represent evolution within family Cladorhizidae rather than evolution of carnivory in separate lineages. Conflicting spicule characters can be reinterpreted to support the inclusion of all carnivorous sponges within Cladorhizidae, and a carnivorous habit should thus be considered the main diagnostic character in systematic classification.

New records of demosponges (Porifera) from the South Orkney Islands (Antarctica) with a checklist for the region.

The South Orkney Islands (SOI) is a poorly studied Antarctic archipelago located in the Scotia Arc. In this study, we added 25 Demospongiae species to the SOI known sponge richness, which was previously represented by only 16 species. Of these, 22 species represent new records for SOI region, although they were previously recorded from other Antarctic sectors. Samples were collected during two Argentinean Antarctic Research Cruises onboard RV Puerto Deseado. The most frequently collected species were Artemisina apollinis, Iophon gaussi, I. unicorne, Lissodendoryx (E.) ramilobosa, Myxodoryx hanitschi and Tedania (T.) tantula. Data and illustrations of some rare or uncommon species are provided: Iophon gaussi, Lissodendoryx (E.) anacantha, Microxina charcoti, Raspailia (H.) hentscheli, Haliclona spongiosissima, Haliclona (G.) cf. cucurbitiformis. Although the present results increase substantially the previously known information on sponge species, this is still an underestimation of total richness because certain orders (Tetractinellida, Suberitida, Biemnida, Polymastiida) and classes were not studied during the current investigation, although previous records were included.

Bone-eating worms from the Antarctic: the contrasting fate of whale and wood remains on the Southern Ocean seafloor.

We report the results from the first experimental study of the fate of whale and wood remains on the Antarctic seafloor. Using a baited free-vehicle lander design, we show that whale-falls in the Antarctic are heavily infested by at least two new species of bone-eating worm, Osedax antarcticus sp. nov. and Osedax deceptionensis sp. nov. In stark contrast, wood remains are remarkably well preserved with the absence of typical wood-eating fauna such as the xylophagainid bivalves. The combined whale-fall and wood-fall experiment provides support to the hypothesis that the Antarctic circumpolar current is a barrier to the larvae of deep-water species that are broadly distributed in other ocean basins. Since humans first started exploring the Antarctic, wood has been deposited on the seafloor in the form of shipwrecks and waste; our data suggest that this anthropogenic wood may be exceptionally well preserved. Alongside the new species descriptions, we conducted a comprehensive phylogenetic analyses of Osedax, suggesting the clade is most closely related to the frenulate tubeworms, not the vestimentiferans as previous reported.

Chemo-ecological studies on hexactinellid sponges from the Southern Ocean.

Hexactinellids (glass sponges) are an understudied class with syncytial organization and poor procariotic associations, thought to lack defensive secondary metabolites. Poriferans, though, are outstanding sources of bioactive compounds; nonetheless, a growing suspicion suggests that many of these chemicals could be symbiont-derived. In Polar latitudes, sponges are readily invaded by diatoms, which could provide natural products. Hexactinellids are typical of deep waters; but in Antarctica, they dominate the upper shelf providing shelter and food supply to many opportunistic mesograzers and macroinvertebrates, which exert strong ecological pressures on them. Aiming to examine the incidence of defensive activities of hexactinellids against consumption, feeding experiments were conducted using their lipophilic fractions. Antarctic hexactinellid and demosponge extracts were tested against the asteroid Odontaster validus and the amphipod Cheirimedon femoratus as putative sympatric, omnivorous consumers. Hexactinellids yielded greater unpalatable activities towards the amphipod, while no apparent allocation of lipophilic defenses was noted. After chemical analyses on the lipophilic fractions from these Antarctic glass sponges, quite similar profiles were revealed, and no peculiar secondary metabolites, comparable to those characterizing other poriferans, were found. Instead, the lipidic compounds 5α(H)-cholestan-3-one and two glycoceramides were isolated for their particular outspread presence in our samples. The isolated compounds were further assessed in asteroid feeding assays, and their occurrence was evaluated for chemotaxonomical purposes in all the Antarctic samples as well as in glass sponges from other latitudes by NMR and MS. Characteristic sphingolipids are proposed as chemical markers in Hexactinellida, with possible contributions to the classification of this unsettled class.

Biodiversity, environmental drivers, and sustainability of the global deep-sea sponge microbiome.

In the deep ocean symbioses between microbes and invertebrates are emerging as key drivers of ecosystem health and services. We present a large-scale analysis of microbial diversity in deep-sea sponges (Porifera) from scales of sponge individuals to ocean basins, covering 52 locations, 1077 host individuals translating into 169 sponge species (including understudied glass sponges), and 469 reference samples, collected anew during 21 ship-based expeditions. We demonstrate the impacts of the sponge microbial abundance status, geographic distance, sponge phylogeny, and the physical-biogeochemical environment as drivers of microbiome composition, in descending order of relevance. Our study further discloses that fundamental concepts of sponge microbiology apply robustly to sponges from the deep-sea across distances of >10,000 km. Deep-sea sponge microbiomes are less complex, yet more heterogeneous, than their shallow-water counterparts. Our analysis underscores the uniqueness of each deep-sea sponge ground based on which we provide critical knowledge for conservation of these vulnerable ecosystems.

Unique spicules may confound species differentiation: taxonomy and biogeography of Melonanchora Carter, 1874 and two new related genera (Myxillidae: Poecilosclerida) from the Okhotsk Sea.

Sponges are amongst the most difficult benthic taxa to properly identify, which has led to a prevalence of cryptic species in several sponge genera, especially in those with simple skeletons. This is particularly true for sponges living in remote or hardly accessible environments, such as the deep-sea, as the inaccessibility of their habitat and the lack of accurate descriptions usually leads to misclassifications. However, species can also remain hidden even when they belong to genera that have particularly characteristic features. In these cases, researchers inevitably pay attention to these peculiar features, sometimes disregarding small differences in the other "typical" spicules. The genus Melonanchora Carter, 1874, is among those well suited for a revision, as their representatives possess a unique type of spicule (spherancorae). After a thorough review of the material available for this genus from several institutions, four new species of Melonanchora, M. tumultuosa sp. nov., M. insulsa sp. nov., M. intermedia sp. nov. and M. maeli sp. nov. are formally described from different localities across the Atlanto-Mediterranean region. Additionally, all Melonanchora from the Okhotsk Sea and nearby areas are reassigned to other genera; Melonanchora kobjakovae is transferred to Myxilla (Burtonanchora) while two new genera, Hanstoreia gen. nov. and Arhythmata gen. nov. are created to accommodate Melonanchora globogilva and Melonanchora tetradedritifera, respectively. Hanstoreia gen. nov. is closest to Melonanchora, whereas Arhythmata gen. nov., is closer to Stelodoryx, which is most likely polyphyletic and in need of revision.

Rock sponges (lithistid Demospongiae) of the Northeast Atlantic seamounts, with description of ten new species.

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 NEA and Mediterranean Sea from 26 to 36 species.

Evidence of Vent-Adaptation in Sponges Living at the Periphery of Hydrothermal Vent Environments: Ecological and Evolutionary Implications.

The peripheral areas of deep-sea hydrothermal vents are often inhabited by an assemblage of animals distinct to those living close to vent chimneys. For many such taxa, it is considered that peak abundances in the vent periphery relate to the availability of hard substrate as well as the increased concentrations of organic matter generated at vents, compared to background areas. However, the peripheries of vents are less well-studied than the assemblages of vent-endemic taxa, and the mechanisms through which peripheral fauna may benefit from vent environments are generally unknown. Understanding this is crucial for evaluating the sphere of influence of hydrothermal vents and managing the impacts of future human activity within these environments, as well as offering insights into the processes of metazoan adaptation to vents. In this study, we explored the evolutionary histories, microbiomes and nutritional sources of two distantly-related sponge types living at the periphery of active hydrothermal vents in two different geological settings (Cladorhiza from the E2 vent site on the East Scotia Ridge, Southern Ocean, and Spinularia from the Endeavour vent site on the Juan de Fuca Ridge, North-East Pacific) to examine their relationship to nearby venting. Our results uncovered a close sister relationship between the majority of our E2 Cladorhiza specimens and the species Cladorhiza methanophila, known to harbor and obtain nutrition from methanotrophic symbionts at cold seeps. Our microbiome analyses demonstrated that both E2 Cladorhiza and Endeavour Spinularia sp. are associated with putative chemosynthetic Gammaproteobacteria, including Thioglobaceae (present in both sponge types) and Methylomonaceae (present in Spinularia sp.). These bacteria are closely related to chemoautotrophic symbionts of bathymodiolin mussels. Both vent-peripheral sponges demonstrate carbon and nitrogen isotopic signatures consistent with contributions to nutrition from chemosynthesis. This study expands the number of known associations between metazoans and potentially chemosynthetic Gammaproteobacteria, indicating that they can be incredibly widespread and also occur away from the immediate vicinity of chemosynthetic environments in the vent-periphery, where these sponges may be adapted to benefit from dispersed vent fluids.

Abyssocladia vaceleti (Porifera, Cladorhizidae): a new deep-sea carnivorous sponge from Patagonia.

This study describes a new species of carnivorous sponge (Family Cladorhizidae) collected in Patagonia, SW Atlantic, off Argentinean waters and the North of the Falkland Islands (Malvinas). The species described here, belongs to the genus Abyssocladia and was collected by dredging and trawling during IEO (Spanish Institute of Oceanography) cruises in the South West Atlantic Ocean from 2007 to 2010 under the Atlantis Project. Abyssocladia vaceleti sp. nov. is characterised by the possession of a long peduncle and flat body with bilaterally symmetrical and apical filaments with a skeleton of tornotes (often polytylotes), styles, abyssochelae, arcuate chelae, sigmancistras and acanthotylostrongyles. This species lives at depths of 901-1547 m.

New deep Cheilostomata (Bryozoa) species from the Southwestern Atlantic: shedding light in the dark.

The present paper reports on 22 bryozoan species collected from 25 localities on the Southwestern Atlantic continental shelf and slope by the Instituto Español de Oceanografía vessel RV Miguel Oliver (2008-2010). Two new genera, Amynaskolia n. gen. and Biconcavus n. gen., and twenty new cheilostome species are described: Amastigia zigzag n. sp., Membranicellaria balanyai n. sp., Figularia dimorpha n. sp., Biconcavus batmani n. sp., Smittina acicularis n. sp., Smittoidea granulosa n. sp., Amynaskolia foramina n. sp., Mawatarius avilae n. sp., Mucropetraliella reticulata n. sp., Ipsibuffonella umbonata n. sp., Microporella gappai n. sp., Fenestrulina curviscutum n. sp., Fenestrulina multiflorum n. sp., Malakosaria cecilioi n. sp., Osthimosia incisa n. sp., Spigaleos simplex n. sp., Reteporella kuklinskii n. sp., Reteporella maryae n. sp., Reteporella taylori n. sp., Orthoporidra nova n. sp.. New taxonomic remarks are also included for two little-known species: Melicerita atlantica Busk, 1884 and Arachnopusia tubula Hayward Thorpe, 1988. Membranicellaria balanyai n. sp., Mawatarius avilae n. sp. and Ipsibuffonella umbonata n. sp. represent the third described species of their genera, whereas Spigaleos simplex n. sp., Malakosaria cecilioi n. sp. and Orthoporidra nova n. sp. are the fourth, fifth and sixth described species of their genera respectively. Spigaleos (Celleporidae) and Arachnopusia tubula (Arachnopusiidae), previously reported only from Antarctica, Ipsibuffonella (Buffonellodidae), reported from tropical and subtropical regions, and Mucropetraliella (Petraliidae) all represent first records for the Patagonian region, expanding their geographic distributions northwards and southwards, respectively. These results highlight the importance of sampling the slope and deep waters of South American and Antarctic margins and basins, filling significant gaps in the knowledge of the biodiversity and biogeographic ranges of certain taxa in these unique and varied ecosystems.

Sponge grounds of Artemisina (Porifera, Demospongiae) in the Iberian Peninsula, ecological characterization by ROV techniques.

Artemisina Vosmaer, 1885 is a poecilosclerid microcionoid sponge genus with 20 valid species, seven of which have been recorded in the Atlantic Ocean.The present study describes Artemisina sponge grounds in Iberia Peninsula. A. transiens is a sponge described in 1890 by Topsent in Galicia (Spain); A. hispanica was also collected in the north of Spain by Ferrer-Hernández (1917); World Porifera Database (WPD) considers at the moment both mushroom-shaped species as synonyms (van Soest et al., 2018), but we have only been able to check the types of A. hispanica. The studied samples were collected in Somos Llungo station and they correspond clearly to those described as A. hispanica by Ferrer-Hernández (1917) and it presents differences in the skeleton with respet to description of A. transiens in the literture. There are no more records after 1917 and there are no data of ecological characterisation nor is there a detailed description of its skeletal composition with Scanning Electron Microscopy. In the previous records the formation of sponge grounds of these species was not known.                                                                                                                          Oceana, the largest international organization focused solely on protecting the world's oceans, has recorded the habitat of Artemisina in Atlantic and Cantabrian waters during a series of ROV cruises for the identification of marine areas with high ecological value that need protection. Its life conditions and associated fauna are described from direct observations for the first time.

Insights into the reproduction of some Antarctic dendroceratid, poecilosclerid, and haplosclerid demosponges.

Sponges are a dominant element of the Antarctic benthic communities, posing both high species richness and large population densities. Despite their importance in Antarctic ecosystems, very little is known about their reproductive patterns and strategies. In our study, we surveyed the tissue of six different species for reproductive elements, namely, Dendrilla antarctica Topsent, 1905 (order Dendroceratida), Phorbas areolatus (Thiele, 1905), Kirkpatrickia variolosa (Kirkpatrick, 1907), and Isodictya kerguelenensis (Ridley & Dendy, 1886) (order Poecilosclerida), and Hemigellius pilosus (Kirkpatrick, 1907) and Haliclona penicillata (Topsent, 1908) (Haplosclerida). Samples of these six species containing various reproductive elements were collected in Deception Island and were processed for both light and transmission electron microscopy (TEM). Even though we were not able to monitor the entire reproductive cycle, due to time and meteorological conditions, we report important aspects of the reproduction of these species. This includes oocyte and embryo morphology and cell ultrastructure, follicular structures and nurse cell activity, as well as vitellogenesis. All species were brooding their embryos within their mesohyl. Both oocytes and embryos were registered in the majority of the studied species, and a single sperm cell being carried to an egg for fertilization was observed in H. penicillata. While the reproductive periods of all species coincided temporally, some of them seemed to rely on a single spawning event, this being suggested by the synchronic oogenesis and embryogenesis occurrence of D. antarctica, P. areolatus and I. kerguelenensis. In contrast, K. variolosa had an asynchronous embryo development, which suggests several larval release events. Our results suggest that differences in the reproductive strategies and morphological traits might succeed in the coexistence of these species at the same habitat avoiding the direct competition between them.

Bone-Eating Worms Spread: Insights into Shallow-Water Osedax (Annelida, Siboglinidae) from Antarctic, Subantarctic, and Mediterranean Waters.

Osedax, commonly known as bone-eating worms, are unusual marine annelids belonging to Siboglinidae and represent a remarkable example of evolutionary adaptation to a specialized habitat, namely sunken vertebrate bones. Usually, females of these animals live anchored inside bone owing to a ramified root system from an ovisac, and obtain nutrition via symbiosis with Oceanospirillales gamma-proteobacteria. Since their discovery, 26 Osedax operational taxonomic units (OTUs) have been reported from a wide bathymetric range in the Pacific, the North Atlantic, and the Southern Ocean. Using experimentally deployed and naturally occurring bones we report here the presence of Osedax deceptionensis at very shallow-waters in Deception Island (type locality; Antarctica) and at moderate depths near South Georgia Island (Subantarctic). We present molecular evidence in a new phylogenetic analysis based on five concatenated genes (28S rDNA, Histone H3, 18S rDNA, 16S rDNA, and cytochrome c oxidase I-COI-), using Maximum Likelihood and Bayesian inference, supporting the placement of O. deceptionensis as a separate lineage (Clade VI) although its position still remains uncertain. This phylogenetic analysis includes a new unnamed species (O. 'mediterranea') recently discovered in the shallow-water Mediterranean Sea belonging to Osedax Clade I. A timeframe of the diversification of Osedax inferred using a Bayesian framework further suggests that Osedax diverged from other siboglinids during the Middle Cretaceous (ca. 108 Ma) and also indicates that the most recent common ancestor of Osedax extant lineages dates to the Late Cretaceous (ca. 74.8 Ma) concomitantly with large marine reptiles and teleost fishes. We also provide a phylogenetic framework that assigns newly-sequenced Osedax endosymbionts of O. deceptionensis and O. 'mediterranea' to ribospecies Rs1. Molecular analysis for O. deceptionensis also includes a COI-based haplotype network indicating that individuals from Deception Island and the South Georgia Island (ca. 1,600 km apart) are clearly the same species, confirming the well-developed dispersal capabilities reported in other congeneric taxa. In addition, we include a complete description of living features and morphological characters (including scanning and transmission electron microscopy) of O. deceptionensis, a species originally described from a single mature female, and compare it to information available for other congeneric OTUs.

Antarctic Porifera database from the Spanish benthic expeditions.

THE INFORMATION ABOUT THE SPONGES IN THIS DATASET IS DERIVED FROM THE SAMPLES COLLECTED DURING FIVE SPANISH ANTARCTIC EXPEDITIONS: Bentart 94, Bentart 95, Gebrap 96, Ciemar 99/00 and Bentart 2003. Samples were collected in the Antarctic Peninsula and Bellingshausen Sea at depths ranging from 4 to 2044 m using various sampling gears. The Antarctic Porifera database from the Spanish benthic expeditions is unique as it provides information for an under-explored region of the Southern Ocean (Bellingshausen Sea). It fills an information gap on Antarctic deep-sea sponges, for which there were previously very few data. This phylum is an important part of the Antarctic biota and plays a key role in the structure of the Antarctic marine benthic community due to its considerable diversity and predominance in different areas. It is often a dominant component of Southern Ocean benthic communities. The quality of the data was controlled very thoroughly with GPS systems onboard the R/V Hesperides and by checking the data against the World Porifera Database (which is part of the World Register of Marine Species, WoRMS). The data are therefore fit for completing checklists, inclusion in biodiversity pattern analysis and niche modelling. The authors can be contacted if any additional information is needed before carrying out detailed biodiversity or biogeographic studies. The dataset currently contains 767 occurrence data items that have been checked for systematic reliability. This database is not yet complete and the collection is growing. Specimens are stored in the author's collection at the Spanish Institute of Oceanography (IEO) in the city of Gijón (Spain). The data are available in GBIF.