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Porifera: Demospongiae: Axinellida: Raspailiidae) from Deep Seamounts of the Western Pacific

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Porifera: Demospongiae: Axinellida: Raspailiidae) from Deep Seamounts of the Western Pacific Zootaxa 4410 (2): 379–386 ISSN 1175-5326 (print edition) http://www.mapress.com/j/zt/ Article ZOOTAXA Copyright © 2018 Magnolia Press ISSN 1175-5334 (online edition) https://doi.org/10.11646/zootaxa.4410.2.7 http://zoobank.org/urn:lsid:zoobank.org:pub:14C3B3C0-2DF9-4893-998F-23D96450C1FD A new species of the sponge Raspailia (Raspaxilla) (Porifera: Demospongiae: Axinellida: Raspailiidae) from deep seamounts of the Western Pacific MERRICK EKINS1,2,6, CÉCILE DEBITUS3, DIRK ERPENBECK4 & JOHN N.A. HOOPER1,5 1Queensland Museum, PO Box 3300, South Brisbane 4101, Brisbane, Queensland, Australia 2School of Biological Sciences, University of Queensland, St Lucia, Queensland, 4072 Australia 3LEMAR, IRD, CNRS, IFREMER, UBO, IUEM, rue Dumont d’Urville, 29280 Plouzané, France 4Dept. of Earth and Environmental Sciences and GeoBio-Center, Ludwig-Maximilians-Universität München, Richard-Wagner-Straße 10, 80333 München, Germany 5Griffith Institute for Drug Discovery, Griffith University, Brisbane 4111, Queensland, Australia 6Corresponding Author. E-mail: [email protected] Abstract A new species of Raspailia (Raspaxilla) frondosa sp. nov. is described from the deep seamounts of the Norfolk and New Caledonia Ridges. The morphology of the species resembles that of a frond or a fern, and its unique highly compressed axial skeleton of interlaced spongin fibres without spicules in combination with a radial extra axial skeleton of a perpen- dicular palisade of spicules, differentiate it from all other species of the subgenus. This species is compared morphologi- cally to all 18 other valid species described in Raspailia (Raspaxilla). Key words: Raspailia, Raspaxilla, Aulospongus, Seamounts, Norfolk, New Caledonia, rhabdostyles Introduction The subgenus Raspailia (Raspaxilla) Topsent, 1913 was most recently reviewed by Hooper (2002), noting that it differed from typical Raspailia in having rhabdose echinating acanthostyles, and from other rhabdose raspailiids, notably Aulospongus Norman, 1878, in having echinating rhabdostyles geometrically very different from the usually longer choanosomal styles (the latter without any basal rhabd). There are currently 18 species recognised (Van Soest et al. 2017), distributed in the Southwest Pacific Ocean (New Caledonia, New Zealand, Torres Strait, northern Great Barrier Reef, Shoalhaven Bight New South Wales), Northwest Pacific Ocean (Japan, Micronesia), Northeast Indian Ocean (Northwest Shelf), Northeast Pacific Ocean (Mexico, Gulf of California), Southeast Pacific Ocean (Galapagos Islands), Northwest Atlantic (North Carolina), Southwest Atlantic Ocean (Malvinas/ Falkland Islands, Southeast Brazil), and Southwestern Indian Ocean subantarctic region (Prince Edward Islands). Bathymetric distributions of species range from the shallow subtidal waters to depths of over 400 metres. The present study describes a new species from deeper waters (200–310m depth) on the New Caledonian and Norfolk Ridges off New Caledonia, and compares it with other species in the subgenus Raspaxilla. The marine biota of this whole region is notably highly diverse (e.g. Payri & Richer de Forges 2007), and the deeper water (bathyal) sponges on these seamounts are otherwise well-documented (e.g. Lėvi & Lėvi 1983, Lévi 1993, Schlacher-Hoenlinger et al. 2005, Ekins et al. 2016). Materials and methods Collection of material. Sponges were collected by dredge from 200 to 310m on the top of deep seamounts off New Caledonia, on the Norfolk and New Caledonia Ridges. The sponges were preserved in ethanol after collection, and transferred to the Queensland Museum. All specimens were registered at the Queensland Museum (QM), and the holotype deposited in the Muséum National d’Histoire Naturelle (MNHN). Accepted by N. de Voogd: 12 Feb. 2018; published: 17 Apr. 2018 379 Morphological analysis. Tissue preparations for light microscopy were made by cutting thin sections through the ectosome and choanosome with a scalpel. These sections were then cleared in a xylene-phenol saturated solution for 12 hours, dried, mounted on a microscope slide and fixed in Durcapan (Sigma Aldrich co., St. Louis, USA). Scanning Electron Microscope spicule preparations were made by dissolving the tissue in 12.5% sodium hypochlorite to remove the soft tissue, and neutralized in distilled water, rinsed twice in 70% ethanol and then finally in 98% ethanol and then air dried. SEM preparations were sputter coated in gold to improve resolution. Some of the SEM tissue preparations were simply dissected and then air dried before gold sputter coating. The scanning electron micrograph photos and measurements were made using a Hitachi TM-1000 SEM and plates assembled in Adobe Photoshop. The spicule size ranges are from those spicules that were the largest and smallest of all spicules from all three specimens. Molecular analysis. DNA was extracted with the DNEasy Mini Kit (Qiagen, Hilden) and with classical Phenol / Chloroform extractions (e.g. Sambrook 1989). PCR reactions for two independent markers (CO1, and C- region of 28S rDNA) were attempted following the procedures as described in Erpenbeck et al. 2016. Results Class Demospongiae Sollas, 1885 Subclass Heteroscleromorpha Cárdenas, Pérez & Boury-Esnault, 2012 Order Axinellida Lévi, 1953 Family Raspailiidae Hentschel, 1923 Genus Raspailia Nardo, 1833 Raspailiidae with a more-or-less compressed axial skeleton and a radial, plumose or simply reticulate extra-axial skeleton, with choanosomal spicules consisting of two, three or more different size classes (styles and/or oxeas), and echinating acanthostyles microcionid-like or secondarily modified (from Hooper et al. 2008). Subgenus Raspaxilla Topsent, 1913 Synonymy: Raspaxilla Topsent, 1913: 616 (Type species: Raspaxilla phakellina Topsent, 1913: 617, by monotypy). Echinaxia Hallmann, 1916a: 543; 1917: 391 (Type species: Axinella frondula Whitelegge, 1907: 509, by original designation). Axinectya Hallmann, 1917: 393 (Type species: Axinella mariana Ridley & Dendy, 1886: 480, by original designation). Definition. Raspailia with echinating rhabdostyles geometrically very different from the usually longer choanosomal styles (the latter without any basal rhabd); extra-axial styles forming a radial skeleton perpendicular to the axis; and well differentiated axial and extra-axial skeletons (the former compressed, the latter plumo- reticulate and/or radial) (from Hooper et al. 2008). Raspailia (Raspaxilla) frondosa sp. nov. (Figs 1–4, Table 1) Etymology. Latin for leafy, reflecting the gross morphology of the sponge resembling the frond of a fern. Material examined. Holotype: MNHN-IP-2015-1402 (fragment of the holotype QM G335124) Station DW3110, New Caledonia Ridge 23o1.35’ S 168o16.99’ E to 23o1.71’S 168o15.86E, 270–310 m, Coll. Cécile Debitus, 28.x.2008 (Samadi 2008). Paratypes: QM G335144 (same data as holotype). QM G318701 NORFOLK 1 Station DW1724, Norfolk 380 · Zootaxa 4410 (2) © 2018 Magnolia Press EKINS ET AL. Ridge Seamount Banc.no.1. 23o17.05’S 168o14.28’E, Dredge 200–291 m, Coll. Bertrand Richer de Forges, 27.vi.2001 (Richer de Forges 2001). FIGURE 1. Fixed specimens of R. (R.) frondosa sp. nov., clockwise from the top left: paratype QM G335144; holotype MNHN-IP-2015-1402 (fragment QM G335124); paratype QM G318701. Description. Growth form: A stalked foliaceous, flabellate to reniform sponge growing mainly in one plane but showing some undulations. The blades are between 55 and 65 mm high and 55 to 85 mm wide, with a maximum thickness of 3 mm at the base sloping to less than 1 mm at the apex. It has a circular stalked base with a diameter of 10 mm and small basal holdfast. Colour: Pale cream to orange on deck and beige to brown in ethanol. Texture: Firm and barely compressible, but easy to tear. Surface: Oscules are not visible, however on the surface the absence of parchment-like ectosome over the aquiferous channels resembles leaf veins (this is could also be due to friction or predation). Otherwise the surface is highly hispid and velvety from protruding long styles. Ectosomal skeleton: Membranous, parchment-like, without a specialised ‘raspailiid’ skeleton of spicule- bouquets, but with a thick palisade of subectosomal styles protruding through the ectosome up to 0.5–2 mm. Choanosomal skeleton: With well-differentiated axial and extra-axial skeletons. The axial skeleton is A NEW SPECIES OF THE SPONGE RASPAILIA (RASPAXILLA) Zootaxa 4410 (2) © 2018 Magnolia Press · 381 compressed, lacking spicules, but with heavy interlaced spongin fibres forming a wide core up to 50% of the diameter of the skeleton. Long subectosomal styles and shorter choanosomal rhabdostyles have their bases embedded in the axial spongin skeleton, standing perpendicular to the axis forming a radial extra-axial skeleton, with the subectosomal styles protruding through the ectosome and the choanosomal rhabdostyles supporting the parchment-like ectosome on both dorsal and ventral surfaces of the sponge. FIGURE 2. Spicules of the holotype R. (R.) frondosa sp. nov., MNHN-IP-2015-1402 (fragment QM G335124). A subectosomal style, B terminations of subectosomal style, C–G choanosomal rhabdostyles FIGURE 3. An undigested cross section of the holotype of R. (R.) frondosa sp. nov., MNHN-IP-2015-1402 (fragment QM G335124), showing the choanosomal rhabdostyles supporting the fragile ectosome and the subectosomal styles penetrating the ectosome. Megascleres (Table 1). There are only two categories of megascleres. Choanosomal rhabdostyles are very smooth, moderately thick, with a smooth non-tylote base, varying
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