An integrative systematic framework helps to reconstruct skeletal evolution of glass sponges (Porifera, Hexactinellida).

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.

Four new species of Cladorhizidae (Porifera, Demospongiae, Poecilosclerida) from the Northeast Pacific.

Interest in cladorhizid sponges has grown rapidly in the past 19 years since a unique feeding strategy, carnivory, was described by Vacelet and Boury-Esnault in 1995. Since that time, 31% of the 133 extant cladorhizids have been described. Previously, seven species of cladorhizid sponges were known from the Northeast Pacific. Here we describe four additional species, including two species of Asbestopluma and two species of Cladorhiza. We report on species ranges, habitat, and ecology, including one from a chemosynthetic environment that appears to be using methane-oxidizing bacteria as a nutrient source. In fact, three of the four species described here were found in chemosynthetic habitats. The presence of small crustacean prey was also documented for three of these species. 

New glass sponges (Porifera: Hexactinellida) from deep waters of the central Aleutian Islands, Alaska.

Hexactinellida from deep-water communities of the central Aleutian Islands, Alaska, are described. They were mostly collected by the remotely operated vehicle 'Jason II' from 494­2311 m depths during a 2004 RV 'Roger Revelle' expedition, but one shallow-water species collected with a shrimp trawl from 155 m in the same area is included. The excellent condition of the ROV-collected specimens enabled valuable redescription of some species previously known only from badly damaged specimens. New taxa include one new genus and eight new species in five families. Farreidae consist of two new species, Farrea aleutiana and F. aspondyla. Euretidae consists of only Pinulasma fistulosum n. gen., n. sp. Tretodictyidae include only Tretodictyum amchitkensis n. sp. Euplectellidae consists of only the widespread species Regadrella okinoseana Ijima, reported here over 3,700 km from its closest previously known occurrence. The most diverse family, Rossellidae, consists of Aulosaccus ijimai (Schulze), Aulosaccus schulzei Ijima, Bathydorus sp. (young stage not determinable to species), Caulophacus (Caulophacus) adakensis n. sp., Acanthascus koltuni n. sp., Staurocalyptus psilosus n. sp., Staurocalyptus tylotus n. sp. and Rhabdocalyptus mirabilis Schulze. We present argument for reinstatement of the abolished rossellid subfamily Acanthascinae and return of the subgenera 　Staurocalyptus Ijima and Rhabdocalyptus Schulze to their previous generic status. These fauna provides important complexity to the hard substrate communities that likely serve as nursery areas for the young stages of commercially important fish and crab species, refuge from predation for both young and adult stages, and also as a focal source of prey for juvenile and adult stages of those same species.

Bathydorus laniger and Docosaccus maculatus (Lyssacinosida; Hexactinellida): Two new species of glass sponge from the abyssal eastern North Pacific Ocean.

Two new species of glass sponge were discovered from the abyssal plain 200 km west of the coast of California (Station M). The sponges have similar gross morphology--an unusual plate-like form with basalia stilling the body above soft abyssal sediments. Bathydorus laniger sp. n. differs from its congeners by the presence of dermal and atrial stauractins; it is also supported by smooth hypodermal pentactins and hypoatrial hexactins. Microscleres include oxyhexasters and oxyhemihexasters. Docosaccus maculatus sp. n. contains large hexactins (>1 cm), characteristic of the genus. Megascleres include dermal hexactins, atrial pentactins, and choanosomal hexactins and diactins. Microscleres include oxytipped hemihexasters and floricomes. Several features serve to differentiate this species from its only known congener.

Expanded sampling of New Zealand glass sponges (Porifera: Hexactinellida) provides new insights into biodiversity, chemodiversity, and phylogeny of the class.

Glass sponges (Hexactinellida) constitute important parts of ecosystems on the deep-sea floor worldwide. However, they are still an understudied group in terms of their diversity and systematics. Here, we report on new specimens collected during RV Sonne expedition SO254 to the New Zealand region, which has recently emerged as a biodiversity hotspot for hexactinellids. Examination of the material revealed several species new to science or so far unknown from this area. While formal taxonomic descriptions of a fraction of these were published earlier, we here briefly report on the morphology of the remaining new species and use the collection to greatly expand the molecular phylogeny of the group as established with ribosomal DNA and cytochrome oxidase subunit I markers. In addition, we provide a chemical fingerprinting analysis on a subset of the specimens to investigate if the metabolome of glass sponges contains phylogenetic signal that could be used to supplement morphological and DNA-based approaches.

Rossellid glass sponges (Porifera, Hexactinellida) from New Zealand waters, with description of one new genus and six new species.

New Zealand's surrounding deep waters have become known as a diversity hotspot for glass sponges (Porifera: Hexactinellida) in recent years, and description and collection efforts are continuing. Here we report on eight rossellids (Hexasterophora: Lyssacinosida: Rossellidae) collected during the 2017 RV Sonne cruise SO254 by ROV Kiel 6000 as part of Project PoribacNewZ of the University of Oldenburg, Germany. The material includes six species new to science, two of which are assigned to a so far undescribed genus; we further re-describe two previously known species. The known extant rossellid diversity from the New Zealand region is thus almost doubled, from nine species in five genera to 17 species in eight genera. The specimens described here are only a small fraction of hexactinellids collected on cruise SO254. Unfortunately, the first author passed away while working on this collection, only being able to complete the nine descriptions reported here. The paper concludes with an obituary to him, the world-leading expert on glass sponge taxonomy who will be greatly missed.

Report of Cladorhiza bathycrinoides Koltun (Demospongiae) from North America and a new species of Farrea (Hexactinellida) among sponges from Cordell Bank, California.

A small collection of five Sponges made by E/V Nautilus on Cordell Bank National Marine Sanctuary, California, include the first report of the crinorhyzid Cladorhiza bathycrinoides Koltun off North America and a new species of Hexactinellida, Farrea. cordelli n. sp. The other three sponges in the collection are known to occur off the surrounding North-East Pacific coast, but new geographic or depth distributions are reported for these.

Characterization of deep-sea benthic invertebrate megafauna of the Galapagos Islands.

The deep sea represents the largest and least explored biome on the planet. Despite the iconic status of the Galapagos Islands and being considered one of the most pristine locations on earth, the deep-sea benthic ecosystems of the archipelago are virtually unexplored in comparison to their shallow-water counterparts. In 2015, we embarked on a multi-disciplinary scientific expedition to conduct the first systematic characterization of deep-sea benthic invertebrate communities of the Galapagos, across a range of habitats. We explored seven sites to depths of over 3,300 m using a two-part Remotely Operated Vehicle (ROV) system aboard the E/V Nautilus, and collected 90 biological specimens that were preserved and sent to experts around the world for analysis. Of those, 30 taxa were determined to be undescribed and new to science, including members of five new genera (2 sponges and 3 cnidarians). We also systematically analysed image frame grabs from over 85 h of ROV footage to investigate patterns of species diversity and document the presence of a range of underwater communities between depths of 290 and 3,373 m, including cold-water coral communities, extensive glass sponge and octocoral gardens, and soft-sediment faunal communities. This characterization of Galapagos deep-sea benthic invertebrate megafauna across a range of ecosystems represents a first step to study future changes that may result from anthropogenic impacts to the planet's climate and oceans, and informed the creation of fully protected deep-water areas in the Galapagos Marine Reserve that may help preserve these unique communities in our changing planet.

Re-description and Proposed Reclassification of Myxilla (Burtonanchora) lacunosa Lambe, 1893.

Lambe described Myxilla lacunosa from a specimen from Vancouver Island. Lambe's sponge was subsequently placed in the subgenus Burtonanchora based on Lambe reporting smooth styles. A number of investigators, including the authors of this paper, report some styles with spines in Lambe's sponge. We redescribe Myxilla lacunosa based on re-examination of Lambe's holotype, and propose to place it in subgenus Myxilla based on the presence of spined styles rather than subgenus Burtonanchora which is defined as genus Myxilla with smooth styles.

Four new species of Hexactinellida (Porifera) and a name replacement from the NE Pacific.

Four new species of Hexactinellida are described from the northwest coast of North America. Two northern ones from bottom longline sablefish traps set on Bowie Seamount off northwest Canada are Pinulasma bowiensis and Rhabdocalyptus trichotis. Two southern forms picked by ROV from the wreck of USS Independence off San Francisco, California are Staurocalyptus pamelaturnerae and Hyalascus farallonensis. A fifth specimen from the southern site is considered conspecific with the junior homonym Farrea aculeata Schulze, and allowed renaming of that species as Farrea schulzei. These additions bring the number of Hexactinellida known from the area (30º-90º N) to 62 species.

New taxa and arrangements within the family Latrunculiidae (Demospongiae, Poecilosclerida).

Extensive new collections of latrunculid sponges from British Columbia, the Aleutian Islands, and the Gulf of Alaska, have extended the distributions of known species Latrunculia oparinae Samaai & Krasokhin, 2002, from the Russian Sea of Okhotsk, L. velera Lehnert et al., 2006, from the Aleutian Islands, and L. austini Samaai et al., 2006, from British Columbia. New material has facilitated detailed re-descriptions of these species and in situ images have improved our understanding of their living morphology and ecology. Several new species of Latrunculia were discovered within these collections: sympatric species Latrunculia hamanni sp. nov. and L. oparinae are described here and differentiated from each other by various field characteristics, and L. lincfreesei sp. nov. is described from the Gulf of Alaska. In terms of their higher systematics, L. austini and L. hamanni sp. nov. are recognisable within the subgenus Latrunculia du Bocage, 1869, and L. lincfreesei sp. nov. in subgenus Biannulata Samaai et al., 2006. A third subgenus, Uniannulata subgen. nov., is proposed for L. oparinae, L. velera and several fossil species in southern New Zealand and Western Australian Eocene fossiliferous sediments. In all of these species the subsidiary whorl, apical whorl, and apex of the anisodiscorhabd are fused. Living species of Uniannulata subgen. nov. are thus far restricted to the North Pacific Ocean. A new genus, Bomba gen. nov., is proposed for a rare latrunculid species, Bomba endeavourensis gen. et sp. nov., discovered in deep waters on British Columbia's Endeavour Ridge. Finally, Latrunclava gen. nov. is proposed for species with long, sceptre-like anisoconicorhabds in addition to the typical, smaller anisodiscorhabds; Latrunclava imago gen. et sp. nov. is described from the central Aleutian Islands.

First Lanuginellinae (Porifera, Hexactinellida, Rossellidae) from the NE Pacific and first species of Doconesthes from the Pacific Ocean.

A new sac-shaped hexactinellid collected from western Canada bearing long lateral prostal spicules was first thought to be a typical Rossellinae.  Subsequent examination of its spiculation proved it to have distinctive strobiloplumicomes, typical of the subfamily Lanuginellinae.  Other spicules showed it to be a member of the monospecific genus Doconesthes, known previously only from the North Atlantic Ocean.  The new species described here as Doconesthes dustinchiversi is only the second known species of the genus and the first to be found in the Pacific Ocean.

Rossella podagrosa Kirkpatrick, 1907-A valid species after all.

In this study we provide evidence that the species Rossella podagrosa Kirkpatrick, 1907, commonly considered a synonym of Rossella racovitzae Topsent, 1901, is truly a valid species. We show that it can be clearly distinguished from other species especially when taking into consideration the in situ habitus of the sponge in combination with the spicules. Furthermore we demonstrate the weaknesses in the so far published synonymy concept for the very complicated genus Rossella Carter, 1872. From this we conclude that the best strategy for further analysis of Rossella and establishment of acceptable synonymies will need to be based on detailed examination of the spicules, the holotypes, and in situ habitus. When possible it will be useful to analyze specimens from all Antarctic oceanographic regions.

A review of the Hexactinellida (Porifera) of Chile, with the first record of Caulophacus Schulze, 1885 (Lyssacinosida: Rossellidae) from the Southeastern Pacific Ocean.

All records of the 15 hexactinellid sponge species known to occur off Chile are reviewed, including the first record in the Southeastern Pacific of the genus Caulophacus Schulze, 1885, with the new species Caulophacus chilense sp. n. collected as bycatch in the deep water fisheries of the Patagonian toothfish Dissostichus eleginoides Smitt, 1898 off Caldera (27ºS), Region of Atacama, northern Chile. All Chilean hexactinellid species occur in bathyal to abyssal depths (from 256 up to 4142 m); nine of them are reported for the Sala y Gomez and Nazca Ridges, with one species each in the Juan Fernandez Archipelago and Easter Island. The Chilean hexactinellid fauna is still largely unknown, consisting of only 2.5 % of the known hexactinellid extant species. Further studies and deep water sampling are essential to assess their ecology and distribution, particularly in northern Chile.

Taxonomic review of Hadromerida (Porifera, Demospongiae) from British Columbia, Canada, and adjacent waters, with the description of nine new species.

The history of sponge collecting and systematics in British Columbia is reviewed over the period 1878 to 1966. Recent additions and changes are provided in an on-line species list: www.mareco/org/kml/projects/NEsponges.asp. Hadromerids are the focus of this paper as eight of 19 species in British Columbia are considered new. An additional new species is described from southern California to clarify the status of Tethya californiana in BC. An update is timely for hadromerids in BC as there is new material and renewed interest, while existing descriptions are often inadequate. We describe new species and provide additions to previous descriptions for sponges of the order Hadromerida (Porifera: Demospongiae) in the cold temperate NE Pacific off British Columbia and adjacent waters. We propose one range extension and one new species in Clionaidae; two range extensions and five new species in Polymastiidae; one range extension, two name changes and two new species in Suberitidae; and one new species in Tethyidae. New species include Pione gibraltarensis n.sp., Polymastia piscesae n. sp., Radiella endeavourensis n. sp., Sphaerotylus raphidophora n. sp., Sphaerotylus verenae n. sp., Weberella perlucida n. sp., Prosuberites saanichensis n. sp., Suberites lambei n. sp., and Tethya vacua n. sp..

Two new species in the family Axinellidae (Porifera, Demospongiae) from British Columbia and adjacent waters.

Two new species of Demospongiae are described for British Columbia and adjacent waters in the family Axinellidae, Auletta krautteri sp. n. and Dragmacidon kishinensis sp. n. They represent range extensions for both of these genera. Both are fairly commonly encountered, Auletta krautteri below diving depths (87 to at least 300 m) and Dragmacidon kishinensis in shallow water (intertidal to 30 m). We propose an amended genus diagnosis for Auletta to account for the variability among species in principal spicules that form the ascending tracts to be either oxeas, styles or strongyles rather than just oxeas.

The sponge pump: the role of current induced flow in the design of the sponge body plan.

Sponges are suspension feeders that use flagellated collar-cells (choanocytes) to actively filter a volume of water equivalent to many times their body volume each hour. Flow through sponges is thought to be enhanced by ambient current, which induces a pressure gradient across the sponge wall, but the underlying mechanism is still unknown. Studies of sponge filtration have estimated the energetic cost of pumping to be <1% of its total metabolism implying there is little adaptive value to reducing the cost of pumping by using "passive" flow induced by the ambient current. We quantified the pumping activity and respiration of the glass sponge Aphrocallistes vastus at a 150 m deep reef in situ and in a flow flume; we also modeled the glass sponge filtration system from measurements of the aquiferous system. Excurrent flow from the sponge osculum measured in situ and in the flume were positively correlated (r>0.75) with the ambient current velocity. During short bursts of high ambient current the sponges filtered two-thirds of the total volume of water they processed daily. Our model indicates that the head loss across the sponge collar filter is 10 times higher than previously estimated. The difference is due to the resistance created by a fine protein mesh that lines the collar, which demosponges also have, but was so far overlooked. Applying our model to the in situ measurements indicates that even modest pumping rates require an energetic expenditure of at least 28% of the total in situ respiration. We suggest that due to the high cost of pumping, current-induced flow is highly beneficial but may occur only in thin walled sponges living in high flow environments. Our results call for a new look at the mechanisms underlying current-induced flow and for reevaluation of the cost of biological pumping and its evolutionary role, especially in sponges.

The aquiferous systems of three marine demospongiae.

The aquiferous systems of three common, coastal, marine Demospongiae, Halichondria panicea (Pallas), Haliclona permollis (Bowerbank) and Microciona Prolifera (Ellis and Solander), are analyzed by measurements of cross-sectional areas of conducting elements. The patterns in demosponges of extremely different organizational morphologies are found to be quantitatively similar. The porocyte nature of the ostia is established for all three species. Choanocyte chamber densities range from 1 to 1.8 × 107 chambers ml-1 with 57 to 95 choanocytes per chamber (means). Cross-sectional area of the intervillar space of the choanocyte collars is calculated to be 12 to 56 times the lateral surface area of the specimen. Velocities of water movement through specific elements of the aquiferous system are calculated from cross-sectional area data and measured oscular flow of Haliclona permollis. The calculated Reynolds numbers lie below the critical value and fluid flow is thus considered laminar throughout the aquiferous systems of these sponges.