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

Accepted by M. Klautau: 26 Jun. 2018; published: 31 Aug. 2018 Licensed under a Creative Commons Attribution License http://creativecommons.org/licenses/by/3.0 ZOOTAXA ISSN 1175-5326 (print edition) ISSN 1175-5334 (online edition) Copyright © 2018 Magnolia Press Zootaxa 4466 (1): 095 – 123 http://www.mapress.com/j/zt/ Article 95 https://doi.org/10.11646/zootaxa.4466.1.10 http://zoobank.org/urn:lsid:zoobank.org:pub:24F4B61D-1B0B-42EB-ABED-45766BB53418 Sponge grounds of Artemisina (Porifera, Demospongiae) in the Iberian Peninsula, ecological characterization by ROV techniques PILAR RÍOS 1,2 , RICARDO AGUILAR 3 , ANA TORRIENTE 3 , ARACELI MUÑOZ 4 & JAVIER CRISTOBO 1,2 1 Instituto Español de Oceanografía. Centro Oceanográfico de Gijón, C/ Príncipe de Asturias 70 bis, 33212 Gijón, Asturias, Spain. E-mail: [email protected]; [email protected] 2 Departamento de Zoología y Antropología Física. Universidad de Alcalá de Henares. Madrid, Spain. 3 Oceana. C/ Gran Vía, 59–9. Madrid, Spain. E-mail: [email protected]; [email protected]. 4 Grupo Multidisciplinar de Cartografiado (Tragsatec). Secretaría General de Pesca (SGP), Valentín Beato 6, 28034 Madrid (Spain). E-mail: [email protected] Abstract 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. Key words: Sponge aggregations, sponge associated fauna, ecology, taxonomy, Atlantic Introduction Sponge grounds are reservoirs of biodiversity for the United Nations Environment Programme (UNEP). In these deep-sea habitats, sponges create an environment that supports high biodiversity, provide refuge for fish species, and are a storehouse of novel chemical compounds, some of which are promising for pharmaceutical drug development (Hogg et al. 2010). There is a great need to improve awareness and understanding of sponge grounds, as many of them are not well-defined. A. transiens and A. hispanica are two poorly-known species and were not known to occur in dense fields until now. In 2008, Oceana organized a cruise in Galicia and Cantabrian waters with the aim of identifying areas that should be protected due to their significant ecological value to improve the European network of marine protected areas (MPAs) (Aguilar et al. 2009). This expedition on board the "Oceana's Ranger", a Ketch catamaran 21 m long, carried out 51 dives with an ROV (Remote Operated Vehicle); bringing about more than 88 effective filmed hours. In ten different locations in Galicia and Asturias and in another one in the south of Spain, a sponge was registered with a characteristic habitus named “mushroom sponge” in the expedition. This species was found at different depths (sometimes in high densities) but appeared to prefer waters deeper RÍOS ET AL. 96 · Zootaxa 4466 (1) © 2018 Magnolia Press than 50 m. In order to identify and study it, a special dive with an ROV and divers was prepared to collect samples. As a result of this dive, 10 specimens were collected in the Somos Llungo area to the east of Peñas Cape (Asturias) at 48–50 m. deep; all of them globular pedunculate with apical oscules and an even surface, similar to description of A. hispanica. Material and methods During July and August 2008, Oceana performed an ocean-going expedition off the Atlantic Galician coast and the southern Gulf of Biscay, sailing around 1500 nautical miles (Fig 1). A survey was made with the research boat Ranger, a Ketch catamaran 21 m long and 9.75 m wide. The ROV employed was a Phantom HD 2+2 from Deep Ocean Engineering, equipped with a digital camera of 750 resolution lines. A total 51 ROV video transects between 9 m and 260 m deep, and 54 dives by divers between 0 and -35 m, were completed. ROV transects lasted between 1 and up to 4 hours depending on the location. It examined distances between 0.3 and 1.2 NM. The ROV sailed at 0.2–0.4 knots positioned a few centimetres above the seafloor with the camera slightly tilted downward so as to obtain a vision field of 1.5–2 m wide and a depth image of 4–5 m. The area surveyed per hour was of approx. 700–850 square metres. 10.2-megapixel Reflex photographic cameras and High Definition HDV video cameras were used by the scuba divers, while the Phantom HD2+2 was used by the ROV. After sponges were located by the ROV, samples were taken by the scuba divers at 50 m in position 43º41.085’N–005º47.177’W. 4,000 photographs, as well as 88h09m of ROV recording and 18h21m of divers’ filmed materials were collected. The area of seafloor covered was approx. 70,500 m 2 . 21 out of 54 divers’ dives and 34 out of the 51 ROV’s dives were carried out from the southeastern Gulf of Biscay (Cantabrian Sea) to the Galician coast, where Artemisina findings were concentrated, collecting 7h06m of ROV recording and 5h07m of filmed materials and 1,500 photographs. Only those 34 ROV transects were taken into consideration in this analysis, covering 45,480 m 2 . Artemisina was found during 19 ROV’s transects in 11 different locations 35–126 m deep (Table I). The morphology of the seafloor of the Somos Llungo area was obtained during one cruise onboard R/V Vizconde de Eza in 2010 in the framework of the study of Vulnerable Marine Ecosystems on the INDEMARES Project (Figs. 1A–B). Navigation was provided by a differential GPS system integrated with the ship's central navigation system. Swath bathymetry data were acquired using a multibeam echosounder Simrad EM-300, which works with a transmission frequency between 26 and 34 kHz, swath opening up to 135º and 135 beams per ping. In addition to echosounding information, the backscattered reflectivity of the seafloor imaged by the multibeam transducers was also recorded in digital form. Data were logged and processed with SIS, Neptune and C-Floor software packages, obtaining a grid resolution of 5 m, with full seafloor coverage meeting the International Hydrographic Organization standards for marine hydrographical surveys. Analyses and representation of bathymetric data were performed with ArcGIS software. Other material examined: Artemisina hispanica Ferrer-Hernández MMC 3/C/120 (5 specimens); MMC 3/C/ 184 (1 specimen) and MMC 3/C/199 (2 specimens). In order to study the spicules, the organic matter was digested by nitric acid taken to the boiling point, following the protocols of Rützler (1978) and Cristobo et al. (1993). The data for spicule sizes are based on 25 measurements for each spicule category, comprising minimum, maximum and average lengths in micrometres (µm). Permanent preparations of skeletal architecture for the specimens were done. Spicules were examined with a Jeol 6100 Scanning Electron Microscope. The classification system adopted in this work is that proposed by Hooper (2002) in the Systema Porifera and updated in the World Porifera Database (Van Soest et al. 2018). Zootaxa 4466 (1) © 2018 Magnolia Press · 97 SPONGE GROUNDS OF ARTEMISINA FIGURE 1. A–C. Sample localities of Artemisina showed the dives with ROV in the north of Spain (Atlantic Ocean and Cantabrian Sea). D. Coloured three-dimensional image of the Somos Llungo area (point of view from the south) mapped during the cruise INDEMARES Aviles-0410. RÍOS ET AL. 98 · Zootaxa 4466 (1) © 2018 Magnolia Press TABLE 1. ROV’s ground verification and bionomic surveys in the Iberian Peninsula NW Spain Sites ROV Nº Dives, Time and m2) Position & Depth of sampling transect Position & Depth of Artemisina findings Torre da Aspa (Portugal) 1 37.05,9819‘N 009.07.2322’W (100-120 m) 37.05,9819‘N 009.07.2322’W (100-120 m) Vigo Ría 1, (0h39m), (520 m2) 42º23,694’N – 008º48.554’W & 42º23.675’N – 008º48.523’W (17-18 m) (no findings) Arousa Ría 3, (4h39m), (3,720 m2) 42º30.541’N – 008º59.799’W & 42º30.562’N – 008º59.659’W (19-23 m) 42º36.059’N – 008º52.771’W & 42º36.051’N – 008º52.878’W (9-18 m) 42º33.095’N – 008º56.552’W & 42º32.977’N – 008º56.584’W (9-22 m) (no findings) (no findings) (no findings) Ons Island 1, (1h36m), (1,280 m2) 42º22.880’N – 008º57.622’W & 42º22.753’N – 008º57.903’W (46-58 m) 42º22.803’N -008º57.937’W & 42º22.805’N -008º57.936’W (51-58 m) Salvora Island 1, (3h50m), (2,867 m2) 42º29.146’N – 009º10.148’W & 42º29.844’N – 09º11.077’W (62-110 m) 42º29.587’N – 009º10.648’W & 42.29.706’N – 009º10.762’W (61-79 m) Villar de Fuentes Bank 2, (5h35m), (4,467 m2) 42º46.489’N – 009º20.511’W & 42º46.661’N – 009º20.268’W (82-128 m) 42º45.327’N – 009º17.230’W & 42º46.184’N – 09º16.525’W (56-107 m) 42º46.514’N – 009º20.503’W & 42º46.607’N – 009º20.633'W (105-127 m) 42º45.387’N – 009º17.193’W & 42º45’797’N – 009º16.799’W (65-97 m) Os Meixidos Bank 1, (1h57m), (1,560 m2) 42º44.887’N – 009º11.924’W & 42º44.797’N – 009.09.167’W (13-37 m) 42º44.887’N – 009º11.924’W (35 m) Sisargas Islands 4, (4h39m), (3,720 m2) 43º23.715’N – 008º52.873’W & 43º23.563’N – 008º51.558’W (57-96 m) 43º21.212’N – 008º48.808’W & 43º21.538’N – 008º48.675’W (54-63 m) 43º21.438’N – 008º51.886’W & 43º21.900’N – 08º51.978’W (35-58 m) 43º22.226’N – 008º49.583’W & 43º22.288’N – 008º49.701’W (30-48 m) 43º23.678’N – 008º52.670’W & 43º23.638’N – 008º52.423’W (57-65 m) (no findings) 43º21.590’N – 008º51.921’W & 43º21.876’N – 008º51.969’W (35-55 m) 43º22.226’N – 008º49.583’W & 43º22.290’N – 008º49.668’W (43-47 m) El Cuervo Bank 1, (2h18m), (1,840 m2) 43º23.853’N – 08º54.114’W & 42º23.987’N – 008º53.428’W (101-127 m) (no findings) Bermeo Bank 4, (7h16m), (5,814 m2) 43º41.000’N – 008º15.152’W & 43º41.004’N & 008º15.168’W (36-44 m) 43º41.638’N – 008º15.829’W & 43º41.596’N & 008º16.200’W (84-106 m) 43º41.741’N – 008º12.260’W & 43º40.768’N – 008º15.780’W (22-121 m) 43º41.676’N – 008º16.969’W & 43º41.142’N – 008º17.173’W (85-143 m) (no findings) 43º41.641’N – 008º15.997’W (99 m) 43º41.433’N – 008º15.500’W & 43º41.431’N – 008º15.504’W (71-72 m) 43º41.553’N – 008º17.033’W & 43º41.518’N – 008º17.024’W (110-123 m) Niebla Bank 1, (2h34m), (2,053 m2) 43º48.842’N – 008º03.388’W & 43º48.997’N – 008º04.175'W (38-105 m) 43º48.887’N – 008º03.813’W & 43º48.887’N – 008º04.818'W (50-52 m) Estaca de Bares 1, (1h34m), (1,253 m2) 43º48778’N – 007º42.098’W & 43º48.848’N – 007º41.679’W (72-86 m) 43º48778’N – 007º42.098’W & 43º48.779’N – 007º41.093’W (72-74 m) Vegadeo Shelf 1, (2h34m), (2.053 m2) 43º49.204’N – 007º34.001’W & 43º49.316’N – 007º34.640’W (116-128 m) 43º49.166’N – 007º34.436’W & 43º49.211’N – 007º34.309’W (116-126 m) Estara Bank 1, (2h36m), (2,080 m2) 43º59.595’N – 006º55.143’W & 43.59º874’N – 006º55.263’W (189-253 m) (no findings) ……continued on the next page Zootaxa 4466 (1) © 2018 Magnolia Press · 99 SPONGE GROUNDS OF ARTEMISINA TABLE 1. (Continued) Sites ROV Nº Dives, Time and m2) Position & Depth of sampling transect Position & Depth of Artemisina findings Aviles Canyon 6, (9h33m), (7,640 m2) 43º42.995’N – 006º04.129’W & 43º43.999’N – 006º04.082’W (226-254) 43º42.906’N – 006º03.871’W & 43º43.130’N – 006º03.867’W (195-235 m) 43º42.281’N – 006º04.453’W & 43º42.308’N – 006º04.356’W (182-222 m) 43º44.186’N – 006º04.751’W & 43º41.148’N – 006º04.738'W (219-224 m) 43º43.159’N – 06º11.051’W & 43º43.204’N – 006º10.783’W (215-239 m) 43º43.104’N – 006º03.577’W & 43º43.083’N – 006º03.733’W (201-243 m) (no findings) (no findings) (no findings) (no findings) (no findings) (no findings) Peñas Cape 1, (1h13m), (973 m2) 43º40.910’N – 005º53.316’W & 43º40.945’N – 005º53.226’W (81-89 m) 43º40.919’N & 005º53.262’W (84 m) Somos Llungo Bank 4, (3h54m), (3,120 m2) 43º41.162’N – 005º47.260’W & 43º41.136’N & 005º47.124’W (59-81 m) 43º41.078’N – 005º47.213’W & 43º41.095’N & 005º47.199’W (49-60 m) 43º41.062’N – 005º47.173’W & 43º41.088’N – 005º47.156’W (48-54 m) 43º40.996’N -005º47.289’W & 43º41.022’N – 05º47.596’W (39-70 m) 43º41.167’N – 005º47.140’W & 43º41.141’N – 005º47.126’W (61-78 m) 43º41.078’N – 005º47.213’W (54 m) 43º41.096’N – 005º47.177’W & 43º41.118’N – 005º47.169’W (48-49 m) 43º41.008’N – 005º47.284’W & 43º41.031’N – 05º47.353’W (54-64 m) Llanes Canyon 1, (0h39m), (520 m2) 43º29.927’N – 004º37.938’W & 43º29.892’N & 04º37.933’W (231-244 m) (no findings) Total 34, (57h06m), (45,480 m2) 42º22.753’N - 008º57.903’W & 43º29.892’N & 04º37.933’W (9-254 m) 42º22.803’N -008º57.937’W & 43º41.167’N – 005º47.140’W (35-126) RÍOS ET AL. 100 · Zootaxa 4466 (1) © 2018 Magnolia Press Systematics (Van Soest et al., 2018) Phylum Porifera Grant, 1836 Class Demospongiae Sollas, 1885 Subclass Heteroscleromorpha Cárdenas, Pérez & Boury-Esnault, 2012 Order Poecilosclerida Topsent, 1928 Family Microcionidae Carter, 1875 Subfamily Ophlitaspongiinae de Laubenfels, 1936 Genus Artemisina Vosmaer, 1885 Artemisina transiens Topsent, 1890 Artemisina hispanica Ferrer-Hernandez, 1917 Description of collected specimens Artemisina hispanica Ferrer-Hernandez, 1917 Diagnosis: Globular pedunculate sponge with several apical oscules that are slightly raised. Mushroom appearance with main body ranging from spherical to elliptical shape. The surface is even but finely hispid; it is similar to the surface of Suberitidae species. Consistency is firm. Surface colour is white, beige, orange or cream-whitish. Ectosome is easily visible and clearer than choanosome. Choanosome is brownish and beige in the peduncle area. Size: up to 6 cm high x 5.5 cm in diametre (Fig 2). Skeleton: Bigger styles are in a confused arrangement in the choanosome and they are arranged in bouquets near the ectosome where the small styles are perpendicular to the surface in a dense palisade. The points of styles protrude at the surface in a finely hispid fashion (Fig 3). Spicules: Styles, palmate isochelae, toxas. Megascleres: The ectosomal styles are smaller and clearly differentiated in size from those of the choanosome, they possess microspined heads: 84.92–(124.19)–243.3 X 1.28–(2.36)–3.85 µm; choanosomal styles likewise have microspined heads and others smooth heads: 307.56–(389.43)–487.04 X 3.05–(4.56)–6.3 µm. Microscleres: Palmate isochelae: 18–(20)–22 µm; toxas thin, shallowly curved, with smooth apices: 38.07– (58.67)–11.29 µm long (Fig. 4). Distribution: South European Atlantic Shelf (MEOW) (Marine Ecoregion of the World) (Van Soest et al. 2018). Galicia, Asturias, Santander (Spain) and São. Vicente Cape (Portugal) (Fig 1). Type locality: 43°44'50"N– 8°12'0"W. 135 m deep, Galicia (Topsent, 1890, 1892). In this paper the distribution limit is increased, as well as the bathymetric distribution: 12–143 m. Collection information of previous records and newly collected specimens were archived in the PANGAEA data repository (https://doi.pangaea.de/10.1594/PANGAEA.892371). Remarks. The genus Artemisina was erected by Vosmaer (1885) for the type species Artemisina suberitoides. The taxon has no real distinctive features, although it differs from other Microcionidae in lacking a distinctive choanosomal skeleton or definite sponging fibres, lacking echinating spicules and having a nearly radial ectosomal skeleton. Due to these reasons, in this work, we describe the species using modern techniques like Scanning Electron Microscopy for the skeleton study and also provide many ecological data to characterize the habitat. There are currently 20 valid species (Van Soest et al. 2018). Artemisina transiens was described by Topsent (1890) from Galician waters and the type locality is close (8 Km) to our video transect in Bermeo Bank and similar depths (36–143 m). Zootaxa 4466 (1) © 2018 Magnolia Press · 101 SPONGE GROUNDS OF ARTEMISINA FIGURE 2. A. Oceana Ranger catamaran and Phantom ROV (courtesy Enrique Talledo). B. Original drawing of A. transiens in original description Topsent, 1892. C. A. hispanica, Ferrer Hernandez, 1917. D. Habitus of A. hispanica, fresh specimen of Somos Llungo bank. E. A. hispanica fresh cut specimen of Somos Llungo bank showing some embryos. F–G. Underwater photos of specimens of Artemisina 46 m deep in Golfo Artabro (courtesy of Javier Souto). The specimens collected in this work in the locality of Somos Llungo correspond exactly to the specimens identified by Ferrer-Hernandez (1917) as A. hispanica. We have not been able to review the holotype of A. transiens and in the World Porifera Database (Van Soest et al. 2018), it is considered that they are synonymous species but there is no explanation as to that. Only Burton (1930) points out that: "There are many remarkable points of resemblance between this species and A. transiens, both from neighbouring localities, and I suspect that if the two holotypes could be re-examined and compared, they would be found to be conspecific". Therefore, Burton thinks that they could be the same species but without being demonstrated so far; thereby, we have decided in this work that both could be valid species until holotypes of both species would be compared. The most significant differences according to the descriptions published by both authors refer to the two clearly differentiated categories of styles in A. hispanica, the size of the toxa and the abundance of smooth styles mixed with the spiny ones. Habitat and Ecological Characterization. Up to now, no ecological data have been recorded for this species (Van Soest et al. 2000; 2018). RÍOS ET AL. 102 · Zootaxa 4466 (1) © 2018 Magnolia Press FIGURE 3. A. hispanica skeleton by Scannig Electron Microscopy. A. Thick section showing the ectosome and choanosome. B–C. Skeletal transversal arrangement of ectosome. D–E. Hispid surface of the sponge. Sites ecological descriptions (Fig 1, Table 1 and Table 2) 1) Ons Island Small rocky bank 1.4 NM W off Ons Island at -45/-60 m surrounded by sandy seafloor with ripple marks, many bivalve molluscs, and brachiopod remains. Rocks were widely covered by the corallimorpharian Corynactis viridis, hydrozoans (e.g. Gymnangium montagui) and several Demospongiae (Cliona celata, Haliclona (Reniera) cinerea, Halichondria (Halichondria) panicea, Guitarra solorzanoi, Tedania (Tedania) urgorrii, Clathria (Microciona) atrasanguinea, Phakellia ventilabrum, Axinella dissimilis, A. polypoides and other unidentified). Red algae Corallinaceae of the genus Lithophyllum and Mesophyllum were present in the shallowest parts. The brachiopod Novocrania anomala was very common, especially in the lower part of the rocks. Echinoderms weere well represented by species such as Holothuria (Panningothuria) forskali, Echinus esculentus, Marthasterias glacialis and Echinaster (Echinaster) sepositus. Other species were the anthozoans Caryophyllia (Caryophyllia) Zootaxa 4466 (1) © 2018 Magnolia Press · 103 SPONGE GROUNDS OF ARTEMISINA smithii, Eunicella verrucosa, Leptogorgia sarmentosa, Alcyonium glomeratum, and Cerianthus membranaceus, the annelida Bonellia viridis, the foraminiferan Miniacina miniacea, the bryozoan Reteporella grimaldi and several annelid polychaetes. More common fish were Labrus mixtus, L. bergylta and Trisopterus luscus. Artemisina cf. transiens was mainly found in the biggest and highest rocks, occupying walls with a 30–45º gradient and a NE orientation, although it was also found on the flat tops. It was absent from the small rocks. It created facies of a few decametres or square metres with the highest densities being 25–30 ind/m 2 . FIGURE 4. A. hispanica spicules by Scannig Electron Microscopy. A. Large style. B–C. Head and point of the large style. D– E. Head and point of the small style. F. Small style. G–K. Different morphologies of style heads. L. Toxa. M. Palmate isochela. RÍOS ET AL. 110 · Zootaxa 4466 (1) © 2018 Magnolia Press fascicularis, G. solorzanoi, Artemisina, D. fruticosum, T. (T.) urgorrii, Geodia cf. barretti, Pachymatisma johnstonia, C. reniformis, H. (H.) panicea, H. (R.) cinerea, C. celata, Petrosia (Petrosia) ficiformis, P. cf. crassa, P. boletiformis, P. mamillaris, Dysidea sp. and others. The other most important phylum was the cnidarians, with the anthozoans D. cornigera, L. sarmentosa, E. verrucosa, P. grayi, A. glomeratum, P. anguiconus, P. axinellae, Epizoanthus sp. and C. viridis, the last one being the most abundant species, although E. verrucosa could form dense “forests”. Also important were the hydrozoans H. halecinum, D. alata, G. montagui, A. abietina, Sertularella spp., P. flabellata, Eudendrium sp., Lafoea sp. and Aglaophenia tubulifera. Other species were the echinoderms E. melo, E. esculentus, E. (E.) sepositus, M. glacialis and H. (P.) forskali, the brachiopods N. anomala and T. retusa, the foraminiferan M. miniacea, the bryozoan Cellaria fistulosa, P. fascialis and S. cervicornis, the polychaetes F. implexa, S. vermicularis, S. spallanzanii and other Sabellidae, the fish Aspitrigla cf. obscura, S. cabrilla, D. vulgaris, Mullus barbatus barbatus, L.bergylta, L. mixtus, C. julis, Symphodus sp., C. rupestris, C. exoletus, A. palloni, Scorpaena loppei, Scorpaena sp., L. piscatorius, P. pollachius, T. luscus, Parablennius gattorougine, T. trachurus, some pleuronectiformes and S. canicula, the crustaceans Pagurus sp. and several Mysida indet. and the molluscs O. vulgaris, P. hirundo, Pruvotfolia pselliotes, C. zizyphinum, C. lampas and Euspira nitida. Artemisina hispanicca was widely distributed and occurred at high densities (up to 30–60 ind/m 2 ) on this bank. The most abundant facies were in the biggest rocks with less turbidity and sedimentation. Some specimens found have two stalks. 12) Golfo Artabro The species was recorded in As Laixiñas, Golfo Artabro, off Ría de Ares (43.46005ºN–8.42348ºW 31/07/2011, 46 m) and Bajo del Zorrón, Golfo Artabro, off Ría de Ferrol (43.455ºN–8.3474ºW 08/08/2014, 38 m) Javier Souto (Pers.com.) (Fig 2A–B). Conclusions Artemisina grounds have been found always on rocky beds from 35 to 126 metres deep. It seems to prefer big, sloped rocks with low turbidity and low sedimentation between -50 and -90 m, but it also occurs on small and flat rocks partially or fully covered by sediment down to -126 m. Although it can be fixed on rocks with different slopes, grades and orientation, including vertical walls, overhangs, the entrance of caves, slabs, etc., this sponge is more abundant in areas typically occupied by suspensivorous species like gorgonians. The density of these sponge grounds can reach up to 50–60 ind/m 2 , looking like “mushroom fields” (Figs. 5 and 6). A characteristic of its morphology is that when specimens are found in areas with high degrees of sedimentation or in places where they must compete with other fauna, the shape of the sponge becomes more elongated and the stalk grows taller. The typical community where this species occurs is dominated by Demospongiae. Most common species found along with it are those of its own order Poecilosclerida (e.g. D. fruticosum, T. (T.) urgorrii, G. solorzanoi, C. (M.) atrasanguinea, A. (A.) dichotoma etc.) and those of the order Axinellida (Phakellia spp., Axinella spp. and A. fascicularis). In many places it occupies a transitional zone between the deep-sea coral (D. cornigera) and the cup sponge (P. ventilabrum) fields. Cnidarians are also common, sometimes the sponges share substrate with dense communities of C. viridis or hydrozoans, and even scattered gorgonians. Brachiopods are also highly abundant on rocks occupied by Artemisina but they choose two different niches; while brachiopods show big colonies in the lower part of the rocks, the sponge prefers the upper zone. Acknowledgements To Jean Vacelet in recognition to a long life dedicated to the study of sponges in many and varied fields of research, for his help to authors solving doubts and for his limitless friendship. Zootaxa 4466 (1) © 2018 Magnolia Press · 111 SPONGE GROUNDS OF ARTEMISINA TABLE 2. List of associated flora and fauna of Artemisina grounds found on the Ons Island (1), Salvora Island (2), Villar de Fuentes Bank (3), Os Meixidos Bank (4), Sisargas Islands (5), Bermeo Bank (6), Niebla Bank (7), Estaca de Bares (8), Vegadeo Shelf (9), Peñas Cape (10) Somos Llungo Bank (11). In Golfo Artabro we haven’t any data regarding associated flora and fauna. Depth range is the depth in meters of the shallowest and the deepest record. Taxa not identified to species level or as putative species are marked with an asterisk (*). Taxon 1 2 3 4 5 6 7 8 9 10 11 Depth OCHROPHYTA <30 Cystoseira baccata (S.G.Gmelin) P.C.Silva, 1952 X <30 Desmarestia ligulata (Stackhouse) J.V.Lamouroux, 1813 X <30 Dictyopteris polypodioides (A.P.De Candolle) J.V.Lamouroux, 1809 X X <30 Dictyota dichotoma (Hudson) J.V.Lamouroux, 1809 X <30 Halidrys siliquosa (Linnaeus) Lyngbye, 1819 X <30 Laminaria hyperborean (Gunnerus) Foslie, 1884 X X <30 Laminaria ochroleuca Bachelot de la Pylaie, 1824 X X X X <30-40 Phyllariopsis purpurascens (C.Agardh) E.C.Henry & G.R.South, 1987 X 30-40 Saccorhiza polyschides (Lightfoot) Batters, 1902 X X <30-40 RODHOPHYTA Corallina sp. X <30 Gelidium corneum (Hudson) J.V.Lamouroux, 1813 X <30 Gelidium sp. X 28 Halurus equisetifolius (Lightfoot) Kützing, 1843 X <30 Lithophyllum incrustans Philippi, 1837 X X X 30-40 Lithophyllum byssoides (Lamarck) Foslie, 1900 X 30 Lithophyllum sp. X 45-60 Mesophyllum sp. X X X X 30-70 Neurocaulon sp. X 28 Peyssonnelia sp. X X X 28-40 Plocamium cartilagineum (Linnaeus) P.S.Dixon, 1967 X <30 Sphaerococcus coronopifolius Stackhouse, 1797 X 28 …….continued on the next page RÍOS ET AL. 112 · Zootaxa 4466 (1) © 2018 Magnolia Press TABLE 2. (Continued) Taxon 1 2 3 4 5 6 7 8 9 10 11 Depth FORAMINIFERA Miniacina miniacea (Pallas, 1766) X X X X X X X X 35-125 PORIFERA Adreus fascicularis (Bowerbank,1866) X X X X X 35-89 Amphilectus cf. fucorum X X 35-80 Antho cf. dichotoma X X 80-128 Antho (Antho) dichotoma (Linnaeus, 1767) X X X X 50-89 Antho cf. involvens X 30-60 Axinella cf. rugosa X 72-86 Axinella damicornis (Esper, 1794) X 72-86 Axinella dissimilis (Bowerbank, 1866) X X X X X X 39-125 Axinella flustra Topsent, 1892 X 39-81 Axinella polypoides Schmidt, 1862 X X X X X X X 39-125 *Axinella spp. X X X X 35-89 *Axinellidae indet. X 39-81 Ciocalypta penicillus Bowerbank, 1862 X 72-86 Clathria (Microciona) atrasanguinea (Bowerbank, 1862) X X X X X X <30-125 Clathrina lacunosa (Johnston, 1842) X 35-70 Cliona celata Grant, 1826 X X X X X X X <30-110 Chondrosia reniformis Nardo, 1847 X X <3081 *Demospongiae indet. X 45-60 Desmacidon fruticosum (Montagu, 1814) X X X X X X X X 35-128 Dysidea sp. X 39-81 Geodia cf. barretti X X X 39-125 Geodia sp. X X X X 50-125 …….continued on the next page Zootaxa 4466 (1) © 2018 Magnolia Press · 113 SPONGE GROUNDS OF ARTEMISINA TABLE 2. (Continued) Taxon 1 2 3 4 5 6 7 8 9 10 11 Depth *Geodiidae indet. X 35-70 Guitarra solorzanoi Cristobo, 1998 X X X X X 35-110 Halichondria cf. bowerbanki X X 55-125 Halichondria (Halichondria) panicea (Pallas, 1766) X X X X X X <30-110 Haliclona cf. fistulosa X X X X 50-125 Haliclona (Reniera) cinerea (Grant, 1826) X X X X X 30-81 Haliclona (Rhizoniera) rosea (Bowerbank, 1866) X 55 *Haliclona spp. X X <30-80 Hemimycale columella (Bowerbank, 1874) X X <30-55 Hymedesmia (Hymedesmia) paupertas (Bowerbank, 1866) X X X 35-125 Hymeniacidon perlevis (Montagu, 1814) X <30 Hymeniacidon cf. perlevis X 30-60 Iophon nigricans (Bowerbank, 1858) X 50-80 Laxosuberites sp. X 50-80 Leucosolenia botryoides (Ellis & Solander, 1786) X <30 Microciona cf. laevis X 72-86 *Microcionidae indet. X 39-81 Mycale (Mycale) lingua (Bowerbank, 1866) X 72-86 Myxilla sp. X X X X 30-128 Pachastrella monilifera Schmidt, 1868 X X X X X 35-128 Pachymatisma johnstonia (Bowerbank in Johnston, 1842) X X <30-81 Petrosia cf. crassa X 39-81 Petrosia (Petrosia) ficiformis (Poiret, 1789) X 39-81 Phakellia cf. robusta X X 35-80 Phakellia robusta Bowerbank, 1866 X X 39-89 …….continued on the next page RÍOS ET AL. 114 · Zootaxa 4466 (1) © 2018 Magnolia Press TABLE 2. (Continued) Taxon 1 2 3 4 5 6 7 8 9 10 11 Depth Phakellia sp. X X 55-128 Phakellia ventilabrum (Linnaeus, 1767) X X X X X X X X X X 39-128 Phorbas fictitius (Bowerbank, 1866) X <30 Plakina sp. X X 50-80 Plakortis sp. X X 35-125 Polymastia boletiformis (Lamarck, 1815) X X 39-86 Polymastia mamillaris (Müller, 1806) X 39-81 Polymastia sp. X 50-80 Quasillina cf. brevis X X 60 - 110 Raspailia (Clathriodendron) cf. hispida X 50-80 Spongosorites sp. X 72-86 Suberites cf. carnosus X X 50-125 Sycon sp. X <30 Tedania (Tedania) pilarriosae Cristobo, 2002 X X <30-60 Tedania sp. X X 80-128 Tedania (Tedania) urgorrii Cristobo, 2002 X X X X X X X X X 35-128 Terpios gelatinosus (Bowerbank, 1866) X <30 Thymosia guernei Topsent, 1895 X <30 CNIDARIA Abietinaria abietina (Linnaeus, 1758) X X X <30-110 Acanthogorgia hirsuta Gray, 1857 X X 50-80 Aglaophenia kirchenpaueri (Heller, 1868) X <30 Aglaophenia tubulifera (Hincks, 1861) X 39-81 Aglaophenia sp. X X 30 - 110 Alcyonium digitatum Linnaeus, 1758 X X 30-80 …….continued on the next page Zootaxa 4466 (1) © 2018 Magnolia Press · 115 SPONGE GROUNDS OF ARTEMISINA TABLE 2. (Continued) Taxon 1 2 3 4 5 6 7 8 9 10 11 Depth Alcyonium glomeratum (Hassall, 1843) X X X X X X <30-125 Alcyonium sp. X 116-128 Antipathella subpinnata (Ellis & Solander, 1786) X X 60-125 Antipathes sp. X X 35-70 Balanophyllia (Balanophyllia) regia Gosse, 1853 X <30 Caryophyllia cf. cyathus X 116-128 Caryophyllia (Caryophyllia) smithii Stokes & Broderip, 1828 X X X X 30-89 *Caryophyllia spp. X X X X <30-125 Cerianthus membranaceus (Gmelin, 1791) X X X X 45-128 Corynactis viridis Allman, 1846 X X X X X X X X X <30-125 Dendrophyllia cornigera (Lamarck, 1816) X X X X X X X X X 39-128 Diphasia alata (Hincks, 1855) X X X X 39-110 Diphasia nigra (Pallas, 1766) X X X 50-125 *Diphasia spp. X X X X 35-128 Epizoanthus arenaceus (Delle Chiaje, 1823) X 116-128 Epizoanthus sp. X X X <30-125 Eudendrium sp. X 39-81 Eunicella verrucosa (Pallas, 1766) X X X X X X X X X X <30-125 Gymnangium montagui (Billard, 1912) X X X X X <30-81 Halecium halecinum (Linnaeus, 1758) X X 39-89 *Hydrozoa indet. X X 45-125 Lafoea sp. X X 39-110 Leptogorgia sarmentosa (Esper, 1789) X X X X X X X X 30-125 Nemertesia anteninna (Linnaeus, 1758) X X 35-110 Paramuricea grayi (Johnson, 1861) X X 39-125 …….continued on the next page RÍOS ET AL. 116 · Zootaxa 4466 (1) © 2018 Magnolia Press TABLE 2. (Continued) Taxon 1 2 3 4 5 6 7 8 9 10 11 Depth Paramuricea cf. grayi X X 50-110 Parantipathes hirondelle Molodtsova, 2006 X 116-128 Parazoanthus anguiconus (Norman, 1868) X X X X 39-128 Parazoanthus axinellae (Schmidt, 1862) X X X X 30-86 Polyplumaria flabellata Sars, 1874 X X X X X X 39-128 Savalia savaglia (Bertoloni, 1819) X X X 55-125 *Scleractinia indet. X 80-125 *Sertularella spp. X X X X <30-110 Swiftia pallida Madsen, 1970 X 60 - 110 Tamarisca tamarisca (Linnaeus, 1758) X 60 - 110 Veretillum cynomorium (Pallas, 1766) X 80-125 ANNELIDA Bonellia viridis Rolando, 1822 X X X X X X X X 35-128 Filograna implexa Berkeley, 1835 X X X X 30-86 Lanice conchilega (Pallas, 1766) X X 72-128 *Polychaeta indet. X 45-60 Polydora sp. X 50-80 Protula sp. X 81-89 Sabella spallanzanii (Gmelin, 1791) X X X X <30-81 *Sabellidae indet. X X X 39-125 Salmacina dysteri (Huxley, 1855) X X X <30-125 Serpula vermicularis Linnaeus, 1767 X X X X X 39-128 Spirobranchus triqueter (Linnaeus, 1758) X 116-128 …….continued on the next page Zootaxa 4466 (1) © 2018 Magnolia Press · 117 SPONGE GROUNDS OF ARTEMISINA TABLE 2. (Continued) Taxon 1 2 3 4 5 6 7 8 9 10 11 Depth ARTHROPODA Balanus sp. X 30-60 *Inachus spp. X X 35-128 Galathea strigosa (Linnaeus, 1761) X 116-128 Goneplax rhomboides (Linnaeus, 1758) X 116-128 *Mysida indet. X X X 39-86 Munida rugosa (Fabricius, 1775) X 116-128 Munida sarsi Huus, 1935 X 35-70 Pagurus sp. X 39-81 Palinurus elephas (Fabricius, 1787) X X 50-125 MOLLUSCA Berthellina edwarsii (Vayssière, 1897) X 55 Calliostoma sp. X X X 35-128 Calliostoma zizyphinum (Linnaeus, 1758) X X 39 - 110 Charonia lampas (Linnaeus, 1758) X X X X 39-125 Eledone cirrhosa (Lamarck, 1798) X X 35 - 110 Euspira nitida (Donovan, 1804) X 39-81 Hypselodoris sp. X 72-86 Mytilus galloprovincialis Lamarck, 1819 X 30-60 Simnia spelta (Linnaeus, 1758) X 35-70 Octopus vulgaris Cuvier, 1797 X X X X X X X 35-128 Pruvotfolia pselliotes (Labbé, 1923) X 39-81 Pteria hirundo (Linnaeus, 1758) X X X X X X X X 35-125 Sepia officinalis Linnaeus, 1758 X 116-128 …….continued on the next page RÍOS ET AL. 118 · Zootaxa 4466 (1) © 2018 Magnolia Press TABLE 2. (Continued) Taxon 1 2 3 4 5 6 7 8 9 10 11 Depth PHORONIDA Phoronis hippocrepia Wright, 1856 X <30 BRYOZOA Caberea ellissii (Fleming, 1814) X <30 Cellaria fistulosa (Linnaeus, 1758) X 39-81 Crisia eburnea (Linnaeus, 1758) X <30 Pentapora fascialis (Pallas, 1766) X X X <30-81 Reteporella grimaldi (Jullien, 1903) X X X <30-128 Smittina cervicornis (Pallas, 1766) X X X X <30-86 BRACHIOPODA *Brachiopoda indet. X X 72-89 Novocrania anomala (O.F. Müller, 1776) X X X X X X X X X 39-128 Megerlia truncata (Linnaeus, 1767) X X X 60-125 Terebratulina cf. retusa X 60 - 110 Terebratulina retusa (Linnaeus, 1758) X X 55-81 ECHINODERMATA Anseropoda placenta (Pennant, 1777) X 35-70 Antedon sp. X 116-128 Amphiura sp. X X 50-128 Aslia lefevrei (Barrois, 1882) X 50-80 Astropecten sp. X 116-128 Echinaster (Echinaster) sepositus (Retzius, 1783) X X X X X X X X X 30-128 …….continued on the next page Zootaxa 4466 (1) © 2018 Magnolia Press · 119 SPONGE GROUNDS OF ARTEMISINA TABLE 2. (Continued) Taxon 1 2 3 4 5 6 7 8 9 10 11 Depth Echinus esculentus Linnaeus, 1758 X X X X X X X X X X 30-128 Echinus melo Lamarck, 1816 X X X X X X X X 35-128 Gracilechinus acutus (Lamarck, 1816) X X X X X 30-128 Hacelia cf. attenuata X 116-128 Holothuria (Panningothuria) forskali Delle Chiaje, 1823 X X X X X X X X X X X 30-125 Leptometra celtica (M'Andrew & Barrett, 1857) X X X 35-80 Luidia ciliaris (Philippi, 1837) X X X 50-89 Luidia sarsii Düben & Koren in Düben, 1844 X 35-70 Marthasterias glacialis (Linnaeus, 1758) X X X X X X X X X 30-128 Ophiopholis aculeata (Linnaeus, 1767) X X X 35-128 Ophiothrix fragilis (Abildgaard in O.F. Müller, 1789) X X X X X 35-128 Paracentrotus lividus (Lamarck, 1816) X 30-60 Parastichopus regalis (Cuvier, 1817) X X X X X 35-128 Spatangus purpureus O.F. Müller, 1776 X 35-70 CHORDATA Acantholabrus palloni (Risso, 1810) X X X X X X 35-128 Ammodytes tobianus Linnaeus, 1758 X 35-70 *Aplydium spp X <30 Arnoglossus sp. X 116-128 Arnoglossus thori Kyle, 1913 X >100 Aspitrigla cf. obscura X X 39-81 Botrylloides leachii (Savigny, 1816) X <30 Capros aper (Linnaeus, 1758) X 50-80 Centrolabrus exoletus (Linnaeus, 1758) X X X X X 35-86 …….continued on the next page