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(Demospongiae: Hadromerida) with Ornamented Exotyles Journal of the Marine Biological Association of the United Kingdom, page 1 of 56. # Marine Biological Association of the United Kingdom, 2016. This is an Open Access article, distributed under the terms of the Creative Commons Attribution licence (http://creativecommons.org/licenses/by/4.0/), which permits unrestricted re-use, distribution, and reproduction in any medium, provided the original work is properly cited. doi:10.1017/S0025315416000655 Polymastiidae (Demospongiae: Hadromerida) with ornamented exotyles: a review of morphological affinities and description of a new genus and three new species alexander plotkin1, christine morrow2, elena gerasimova3 and hans tore rapp1,4,5 1Department of Biology, University of Bergen, Postbox 7803, 5020 Bergen, Norway, 2Department of Zoology, Ryan Institute, National University of Ireland Galway, University Road, Galway, Ireland, 3Ra˚dgivende Biologer AS, Bredsga˚rden, Bryggen, 5003 Bergen, Norway, 4Centre for Geobiology, University of Bergen, Postbox 7803, 5020 Bergen, Norway, 5Uni Environment, Uni Research AS, Postbox 7810, 5020 Bergen, Norway 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 rela- tionships between the polymastiid taxa. Keywords: sponges, Demospongiae, Polymastiidae, morphological affinities, new species Submitted 28 January 2016; accepted 13 April 2016 INTRODUCTION species, T. joubini from Azores, which was notable for the denticulate distal ornamentations on its cortical spicules, Sponges of the family Polymastiidae Gray, 1867 have a simple and Sphaerotylus for Vosmaer’s P. capitata, which was charac- spicule assortment which is usually limited to several size terized by the spherical swellings on its projecting spicules. To categories of smooth monactines (Boury-Esnault, 2002). identify these spicules with usual tyles on the proximal However, in addition to these common spicules, some species extremities and ornaments on the distal extremetities protrud- also possess distally ornamented monactines. This additional ing above the sponge surface Topsent used the term exotyle category of spicules was first recorded in polymastiids by introduced by him 2 years earlier (Topsent, 1896) for the Sollas (1882) who noticed the rounded swellings on the similar spicules in Gomphostegia loricata (now Mycale distal tips of projecting monactines in his new species (Rhaphidotheca) loricata, see Van Soest et al., 2015) from Radiella schoenus from the Norwegian coast. Three years the family Mycalidae. later Vosmaer (1885) recorded similar spicules in his new For the time being nine species of Sphaerotylus from species Polymastia capitata from the Arctic. Dendy & Ridley various locations in polar and temperate waters of both hemi- (1886) noted the similarity between R. schoenus and P. capi- spheres, two species of Proteleia from the southern hemi- tata relegating the latter to synonymy with the former. They sphere and two species of Tylexocladus, one from the North also established a new genus, Proteleia, for their new species, Atlantic and the other from the South Pacific are recognized P. sollasi from South Africa, which was distinguished by the as valid (Van Soest et al., 2015). Exotyles have also been grapnel-like distal ornamentations of its protruding spicules. recorded in Trachyteleia stephensi Topsent, 1928 and in two In 1898 Topsent erected two more polymastiid genera dis- New Zealand species of Polymastia Bowerbank, 1864, P. playing ornamented monactines, Tylexocladus for his new tapetum Kelly-Borges & Bergquist, 1997 and P. umbraculum Kelly-Borges & Bergquist, 1997. Affinities between all these taxa have been discussed (Kelly-Borges & Bergquist, 1997; Corresponding author: Boury-Esnault, 2002), but they have never been properly A. Plotkin revised, and there is still no agreement on the differences at Email: [email protected] the generic level. 1 Downloaded from http:/www.cambridge.org/core. Universitetsbiblioteket i Bergen, on 02 Dec 2016 at 07:29:16, subject to the Cambridge Core terms of use, available at http:/www.cambridge.org/core/terms. http://dx.doi.org/10.1017/S0025315416000655 2 alexander plotkin et al. In this paper we review all known species and varieties of S. exotylotus Koltun, 1970 Proteleia, Sphaerotylus, Trachyteleia and Tylexocladus along S. isidis (Thiele, 1905) comb. nov. with those species of Polymastia which display ornamented S. raphidophora Austin, Ott, Reiswig, Romagosa & McDaniel, exotyles. We establish a new genus, Koltunia gen. nov. for 2014 the Antarctic species Proteleia burtoni Koltun, 1964, describe S. renoufi sp. nov. three new species of Sphaerotylus – from South Africa, Ireland S. sceptrum Koltun, 1970 and West Greenland and propose the transfer of two South S. strobilis sp. nov. Pacific species of Polymastia, one to Sphaerotylus, the other S. tjalfei sp. nov. to Proteleia. Finally, we reconsider the affinities of the S. vanhoeffeni Hentschel, 1914 species studied based on multiple morphological characters. S. verenae Austin, Ott, Reiswig, Romagosa & McDaniel, 2014 Genus Trachyteleia Topsent, 1928 T. stephensi Topsent, 1928 MATERIALS AND METHODS Genus Tylexocladus Topsent, 1898 T. hispidus Le´vi, 1993 This study was based on the type specimens and other T. joubini Topsent, 1898 material stored in Ulster Museum, Belfast (BELUM), Natural Incertae sedis History Museum, London (BMNH), Go¨teborg Natural Polymastia umbraculum Kelly-Borges & Bergquist, 1997 History Museum (GNM), Muse´um National d’Histoire Naturelle, Paris (MNHN), Muse´eOce´anographique de Monaco (MOM), Museum of New Zealand, Te Papa Description of taxa Tongarewa, Wellington (NZNM), National Museum of Family POLYMASTIIDAE Gray, 1867 Natural History, Leiden (RMNH), Smithsonian National Museum of Natural History, Washington (USNM), diagnosis Zoological Institute of Russian Academy of Sciences, Saint- Petersburg (ZIN RAS), Museum fu¨r Naturkunde, Berlin Sponges of massive, encrusting, globular, discoid or peduncu- (ZMB), University Museum of Bergen (ZMBN) and Natural late growth form. Surface slightly velvety to very hispid. History Museum of Denmark, University of Copenhagen Choanosomal skeleton composed of radial megasclere tracts. (ZMUC). Additional fresh material was collected from the A complex specialized cortical skeleton is developed to a Norwegian coast during cruises by the University of Bergen. greater or lesser degree, composed of at least a palisade of The architecture of the sponge skeletons was examined under tylostyles, or oxeas and/or exotyles. Spicules comprise two or light microscope on histological sections prepared on a more size categories and include tylostyles, subtylostyles, precise saw with a diamond wafering blade after embedding strongyloxeas, styles or oxeas. Free spicules are always sponge fragments in epoxy resin as described by Boury- present in the choanosome; they may be intermediary or Esnault et al. (2002), Vacelet (2006) and Boury-Esnault & small tylostyles as well as various microscleres including Be´zac (2007). Spicules were examined under light microscope smooth centrotylote microxeas, acanthose microxeas, raphides and SEM after their isolation from organic matter in nitric in trichodragmata and astrotylostyles. A fringe of long spicules acid following standard procedures. The number of specimens is often present bordering the edge of the body where it is in used for spicule measurements is given in the corresponding contact with the substratum (from Plotkin & Janussen, 2008). section of the description of each species. The number of Genus Koltunia gen. nov. spicules of each category measured in one specimen is indicated as N. Measurements are presented as minimum– type species mean–maximum, unless otherwise indicated. Proteleia burtoni Koltun, 1964 (designation herein). SYSTEMATICS diagnosis Thickly encrusting sponges with shaggy surface. Main choa- Systematic index nosomal skeleton composed of longitudinal tracts of large styles and subtylostyles. These tracts ascend forming cortical Class DEMOSPONGIAE Sollas, 1885 bouquets and a thick surface hispidation. Auxiliary choanoso- Suborder HETEROSCLEROMORPHA Ca´rdenas, Perez & mal skeleton comprises free-scattered small tylostyles. Cortex Boury-Esnault, 2012 and surface hispidation reinforced by small tylostyles and Order POLYMASTIIDA Morrow & Ca´rdenas, 2015 giant exotyles (several mm in length). Distal extremities of Family POLYMASTIIDAE Gray, 1867 the exotyles with several long claws resembling grapnels. Genus Koltunia gen. nov. K. burtoni (Koltun, 1964) comb. nov. etymology Genus Proteleia Dendy & Ridley, 1886 Named after the late Dr Vladimir M. Koltun, the greatest P. sollasi
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