Revision of Rhabdastrella Distincta (Thiele, 1900)
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Revision of Rhabdastrella distincta (Thiele, 1900) A re-description of the holotype and description of two recently re-collected specimens Anne Twaalfhoven (10149996) Biology BSc Thesis, July 2016 University of Amsterdam: dr. P. (Peter) Roessingh, [email protected] Naturalis Biodiversity Center, Leiden: dr. N.J. (Nicole) de Voogd, [email protected] Abstract Sponge taxonomy is largely founded upon outdated descriptions originating from pre-modern times. Luckily, the material that these descriptions are based upon is often still available in Natural History Collections. In this article two sponge samples collected in 2009 from Ternate, Indonesia are compared to the type material of Rhabdastrella distincta. The holotype was described in 1896 by Thiele (1900). This article reviews the material for the first time ever since. The two recently collected samples are the first records of the species since its description. Descriptions of all three samples are based upon external morphology, skeletal structure, spicule measurements and spicule analysis by electron microscopy. It is revealed that, contrary to previous descriptions, the spherasters of R. distincta are stubbed with spines. Consequently, it is argued that the relation of R. distincta to its fellow species should be reconsidered. Euasters of R. distincta POR 5341 Twaalfhoven, July 2016, UvA Introduction When researching environmental change and biodiversity, Natural History Collections (NHC’s) can be of high value. Former taxonomists collected and described large collections of specimens that now serve as the foundation of our current knowledge on marine biodiversity (Jackson, 2001; Hooper & van Soest, 2002; Hoeksema et al, 2011). For reef systems, systematic ecological observations and methodological sampling started no earlier than the 1930’s (Jackson, 2001), and most recent surveys only encompass relatively short timespans (Jackson, 2001; van der Meij et al., 2010; Lister, 2011; Hoeksema et al., 2011). Especially for reef systems, NHC’s can serve as historical baselines in monitoring long term biodiversity changes and designating indicator species. NHC’s are thus useful tools in justifying signals of environmental change (Suarez & Tsutsui, 2004; Hoeksema et al., 2011; de Voogd, 2012). One such NHC was obtained by the German professor W.G. Kükenthal in 1896. Amongst other organisms, he collected about 100 specimens of sponges near the island of Ternate in the Moluccan Sea. Kükenthal’s collection was described by Kieschnick (1896), but his findings were later revised by Thiele (1900 & 1903). These works still serve as the most important sponge records for the area. Their papers contained 51 new species, some of which have never been recorded since and most likely are endemic or even singletons (de Voogd, 2012). It wasn’t until 2009 that the same area was sampled again, this time by the combined efforts of Naturalis (The Netherlands) and E-Win, LIPI (Indonesia) (Hoeksema & van der Meij, 2010). In this expedition over 300 sponge specimens were obtained. Of the species found by Kükenthal, a mere 30 were re-collected. These recent samples offer the opportunity to evaluate the descriptions of the German authors. Rhabdastrella distincta is one of these re-collected species. R. distincta was originally described as Coppatias distinctus by Thiele (1900). In 1903 Thiele merged the genera Coppatias and Jaspis. Consequently, all species that lacked small oxeas were placed into the new genus Rhabdastrella, R. distincta becoming the type species for this particular genus and species. R. distincta is now classified within the family Anacorinidae Schmidt 1870 and defined as ‘- an Ancorinidae with euasters, among which large spherasters or sterrospherasters are abundant. Triaenes may be reduced or absent’ (Uriz, 2002). Thiele describes C. distinctus to be overgrown with a Gellius sponge (now Haliclona (Gellius) Gray 1867). Due to this cryptic growth form, the species is extremely hard to find and to our knowledge, there are no records of R. distincta growing without a sponge epibiont. In fact, after its description, the species has never been collected again. Since Thieles short description, the species has never been properly reviewed. During the Ternate expedition in 2009, two specimens were obtained that show strong resemblance to R. distincta. This article compares the newly collected specimens to the type material. Additionally, a re-description of the type material is given, substantiated with electron microscope (SEM) images. Materials & Methods Specimen collection In November 2009, 300 specimens were collected using SCUBA from the reefs of Ternate, in the Northern Mollucan sea. The specimens were preserved in 70% ethanol and deposited in the sponge collection of the Naturalis Biodiversity Centre (RMNH), in the Netherlands. The NHC collected by Küktenthal in 1896 is located in the Natural History Museum of Berlin. Sub samples of the specimens were taken and are kept in Naturalis. Specimen Analysis Specimen descriptions are based on external morphology, skeletal architecture and spicule morphology. For skeletal analysis, tangential sections of the ectosome and perpendicular sections of the choanosome were cut by hand. The sections were air-dried and mounted in Ultrabed on a microscope slide. The slides were studied under a Leitz high power light microscope. Spicule preparations were made by dissolving a small piece of the specimen in 60% Nitric Acid (HNO3) after which the residue was rinsed three times with water and three times with 96% ethanol. The spicules were air-dried on microscopic slides and mounted in Ultrabed. Additionally, spicules were put on aluminium stubs and coated with Argon so that they could be photographed with a Jeol 2 Twaalfhoven, July 2016, UvA Scanning Electron Microscope (SEM). Spicules were measured using the Leitz high power light microscope in combination with Leica Application Suite V4.8. Oxea dimensions are given as a range of length measurements x range of width measurements. For oxyasters and oxyspherasters only the range in diameter is given. For spherasters, ray length was measured as well. Spicule-size ranges are taken from 20-30 measurements. Abbreviations used in this article are RMNH (Naturalis Centre for Biodiversity, Leiden) and ZMB (Museum für Naturkunde und der Universität Humbolt zu Berlin, Berlin, Germany). Classification is done in accordance to Systema Porifera (Hooper & van Soest, 2002). Results Class Desmospongiae Order Astrophorida Family Ancorinidae Schmidt 1870 Genus Rhabdastrella Thiele 1903 Rhabdastrella distincta, Thiele 1903 Material examined: Holotype ZMB. 3190, Indonesia, Ternate, Kükenthal leg., Kütkenthal ded.,3.XII.1902 External morphology: To Thiele, only a 2 x 1 cm piece of the holotype was at hand. To me, only a photograph of the specimen was available (fig.1). The specimen’s colour is black. According to Thiele it was overgrown with Gellius couchi (now accepted as Haliclona (Gellius) fibulata (Schmidt, 1862)). Skeletal analysis: Ectosomal and choanosomal slides are unclear. Arrangement in spicules is hard to find. Oxeas appear to be dispersed confusedly in choanosome. High abundancy of spherasters and oxyasters in ectosome (fig.3d). Spicules: Megascleres: large oxeas, fusiform, 646 -1012 um x 17- 27um (905 x 21, n= 20). Short, pointy ends. Microscleres: eurasters: Oxyasters having 6 rays of equal length. Diameter 62 um – 101 um (82, n=20). Spherasters having 12-16 rays, ray length 6 um – 17 um (11, n=29), diameter 22 um – 59 um (41, n= 29). Centrum takes up about 3/4th of diameter. Rays have spines perpendicular to ray axis getting more abundant near the ends. Oxyspherasters having about 13 rays emerging from small centrum. Diameter 16 um – 26 um (22, n=20). Material examined: RMNH POR. 5341, Indonesia, Halmahera, Maitara Maitara NW, stat TER.010, N 0*44’32”, E 127*21’50.9”,29-10-2009, depth unknown, Fieldnr #TER10/291009/092, coll. N.deVoogd External morphology: Piece is largely overgrown with two other species (fig.2a,b). Color after preservation in alcohol is dark green to black. Some pinkish and white growth/sediment attached. Shape is bulky (fig.2a,b). Structure is very firm and resilient. Leather-like ectosome. Rest of the body crumbles apart when cut into thin slices. Dimensions: 11 x 4 x 1 cm. Skeletal analysis: Ectosomal and choanosomal slides are quite unclear. Confusedly dispersed oxeas in choanosome, spherasters densely concentrated in ectosome. Spicules: Megascleres: large oxeas, fusiform, 684 – 991 um x 12 – 20 um (867 x 16, n=20). Ends are short and pointy. Microscleres: Oxyasters having 6-7 rays of mostly equal lengths. No centrum. Diameter 43 um – 92 um (63, n=21). Spherasters having 8-13 rays. Ray length 5 um – 18 um (11, n=22). Diameter 20 um- 52 um (39, n=22). Rays are smooth with an occasional spine at the tip. Oxyspherasters having about 13 -15 rays, no centrum. Diameter 16 um- 25 um (20, n=21) (fig. 6). Material examined: RMNH POR. 5554, Indonesia, Halmahera, off Tidore Pulau Pilongga S, N 0*42’44.1”, E 127*28’47.3”, 12-11-2009, depth 20m, Fieldnr #TER35/121109/?, coll. ? 3 Twaalfhoven, July 2016, UvA External morphology: Rather thin layer overgrowing another sponge species, layer gets thicker and more like a cushion towards the end. Ectosome feels leather-like, however rough at some spots. Color is dark green to black after preservation in alcohol. Several pinkish and white spots of sediment or colonial growth (fig.2c,d). Dimensions: 8 x 2 x 1 cm (Haliclona included. Over most of the specimen, R. distincta forms a 2 mm thick crust). Skeletal analysis: Dense concentration of spherasters in cortex is clearly visible (fig.3b). Bundles of oxeas in endosome. Oxyasters are scattered between oxeas. Spicules: Megascleres: Large oxeas, fusiform, 842 um – 967 um x 7 um – 23 um (776 x 13, n=24). Tips are pointy but slightly rounded. Oxyasters having 5-6 rays emerging from small centrum. Diameter 76 um - 98 um (84, n= 8). Spherasters having 11-15 rays. Ray length 10 um- 13um.