Exitomelita Sigynae Gen. Et Sp. Nov.: a New Amphipod from the Arctic Loki Castle Vent Field with Potential Gill Ectosymbionts
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Heriot-Watt University Research Gateway Exitomelita sigynae gen. et sp. nov.: a new amphipod from the Arctic Loki Castle vent field with potential gill ectosymbionts Citation for published version: Tandberg, AH, Rapp, HT, Schander, C, Vader, W, Sweetman, AK & Berge, J 2012, 'Exitomelita sigynae gen. et sp. nov.: a new amphipod from the Arctic Loki Castle vent field with potential gill ectosymbionts', Polar Biology, vol. 35, no. 5, pp. 705-716. https://doi.org/10.1007/s00300-011-1115-x Digital Object Identifier (DOI): 10.1007/s00300-011-1115-x Link: Link to publication record in Heriot-Watt Research Portal Document Version: Publisher's PDF, also known as Version of record Published In: Polar Biology General rights Copyright for the publications made accessible via Heriot-Watt Research Portal is retained by the author(s) and / or other copyright owners and it is a condition of accessing these publications that users recognise and abide by the legal requirements associated with these rights. Take down policy Heriot-Watt University has made every reasonable effort to ensure that the content in Heriot-Watt Research Portal complies with UK legislation. If you believe that the public display of this file breaches copyright please contact [email protected] providing details, and we will remove access to the work immediately and investigate your claim. Download date: 24. Sep. 2021 Polar Biol (2012) 35:705–716 DOI 10.1007/s00300-011-1115-x ORIGINAL PAPER Exitomelita sigynae gen. et sp. nov.: a new amphipod from the Arctic Loki Castle vent field with potential gill ectosymbionts Anne Helene Tandberg • Hans Tore Rapp • Christoffer Schander • Wim Vader • Andrew K. Sweetman • Jørgen Berge Received: 18 June 2011 / Revised: 6 October 2011 / Accepted: 10 October 2011 / Published online: 2 November 2011 Ó The Author(s) 2011. This article is published with open access at Springerlink.com Abstract The newly discovered deep-sea hydrothermal Keywords Vent fauna Á Norwegian Sea Á Melitidae Á vent field ‘‘Loki’s Castle’’ at 2,350 m depth at 70°N on the Symbioses Á Arctic mid-ocean ridge Á Hydrothermal vents Á Knipovich Ridge north of the island Jan Mayen is the only Stable isotopes known black smoker field from the Arctic Ridge system. This vent field holds a unique fauna clearly distinct from vent sites along the Mid-Atlantic Ridge south of Iceland. In Introduction addition to numerous maldanid and ampharetid polychae- tes one animal of particular interest at this vent site, is a Hydrothermal vent systems along mid-ocean ridges have new genus and species of melitid amphipod. This new been intensively studied for more than 30 years (e.g., species is found in cracks and crevices on vent chimneys as Ballard 1977; Grassle et al. 1979; Hessler and Lonsdale well as in areas where diffuse venting is common such as 1991; Tunnicliffe 1991; Van Dover et al. 2002; the base of vent chimneys. Here, we present a formal Desbruye`res et al. 2006). While hydrothermal vent systems morphological description of this new melitid species and at fast spreading ridges (e.g., The East Pacific Rise) have hypothesize about the main form of its nutrition using received the most attention so far, hydrothermal ecosys- stable isotope and molecular data. tems at slow and ultra-slow spreading ridges are very poorly studied (Schander et al. 2010). The Loki’s Castle A. H. Tandberg (&) C. Schander Institute of Marine Research, PO Box 1870, Nordnes, University Museum of Bergen, PO Box 7800, 5817 Bergen, Norway 5020 Bergen, Norway e-mail: [email protected] W. Vader A. H. Tandberg Tromsø University Museum, University of Tromsø, International Research Institute of Stavanger, 9037 Tromsø, Norway PO Box 8046, 4068 Stavanger, Norway A. K. Sweetman H. T. Rapp Norwegian Institute for Water Research (NIVA), Department of Biology, University of Bergen, Thormøhlensgate 53D, 5006 Bergen, Norway Thormøhlensgate 53a, 5020 Bergen, Norway J. Berge H. T. Rapp Á C. Schander Á A. K. Sweetman University Centre at Svalbard, PO Box 156, Centre for Geobiology, University of Bergen, 9172 Longyearbyen, Norway Alle´gaten 41, 5020 Bergen, Norway J. Berge H. T. Rapp Faculty for Biosciences, Fisheries and Economics, Uni Environment, Thormøhlensgate 49B, University of Tromsø, 9037 Tromsø, Norway 5020 Bergen, Norway 123 706 Polar Biol (2012) 35:705–716 vent field at 2,350 m depth on the Mohn-Knipovich Ridge the Melitidae s. str., several informal groups have been north of Jan Mayen in the Norwegian Sea was the first recognized based on morphology. The combination of an black smoker vent field to be discovered on an ultra-slow extremely reduced inner ramus and a very long outer ramus spreading ridge and the first along the Arctic ridge system on uropod 3 (U3) distinguishes the Melita, Eriopisa, and (Pedersen et al. 2010). The field comprises a number of Eriopisella groups from other Melitidae groups. Among active black smoker chimneys up to 13 m tall at the top of the most abundant macroorganisms at the Loki’s Castle an extensive mound of hydrothermal sulfide deposits. On Vent Field is a new melitid amphipod species that does not the northeastern flank of the mound lies a sedimentary area belong to any of the previously described genera. Here, we with low-temperature diffuse venting and small barite present a formal description and erect a new genus within chimneys (Pedersen et al. 2010; Kongsrud and Rapp 2011). the Melita group to encompass this new species of vent Earlier studies on shallow vent sites on the ridge system amphipod and present a hypothesis about its main form of north of Iceland and around Jan Mayen have shown that the nutrition based on morphological characters as well as fauna is dominated by shallow water and bathyal species molecular and stable isotope data. from the surrounding waters, with just a few examples of species adapted to live at hydrothermal vent systems (Fricke et al. 1989; Schander et al. 2010). Recent expedi- Materials and methods tions to study the marine biology at Loki’s Castle have revealed a completely different picture of Arctic vent Melitid amphipod specimens from the Loki’s Castle vent ecosystems with the fauna characterized by a high degree field were collected over three research cruises between of endemism and local adaptation (Pedersen et al. 2010; 2008 and 2010 aboard the Norwegian R/V ‘‘G.O. Sars’’ Kongsrud and Rapp 2011) and reveal closer links to Pacific (Fig. 1a; Table 1). Samples were collected using the vent faunas compared to vent sites in the Atlantic south of ‘‘Bathysaurus’’ XL remotely operated vehicle (ROV) from Iceland (Pedersen et al. 2010). Argus Remote Systems. Video images were acquired using The amphipod fauna of hydrothermal vents in the a high-definition (HD) camera, and still photos were sub- Pacific is usually dominated by scavengers such as lysi- sequently captured from the video. Fauna samples were anassoids (Barnard and Ingram 1990; Vinogradov 1995a, taken both from the chimney walls as well as in sedi- b), the swarming pardaliscid Halice hesmonectes Martin mentary areas where diffuse venting was seen and in which et al., 1993, and the uristid Ventiella sulfuris Barnard and dense fields of the siboglinid tube worm Sclerolinum Ingram, 1990 (Van Dover et al. 1992; Martin et al. 1993). contortum were observed. The samples were sorted on In the Atlantic Ocean, the eusirids Bouvierella curtirama board and fixed in either 96% alcohol or 6% buffered Bellan-Santini and Thurston, 1996 and Luckia striki Be- formaldehyde in seawater. Type material has been depos- llan-Santini and Thurston, 1996 are the only abundant and ited in the Natural History Collections, Bergen Museum, swarm-forming amphipods known from hydrothermal University of Bergen, Norway (ZMBN) (Table 1). vents (Van Dover et al. 1992; Martin et al. 1993; Sheader In the laboratory, specimens were dissected in ethanol et al. 2004; Bellan-Santini 2006). and glycerol, and dissected mouth parts, head appendages, Other amphipods reported in lower numbers at Atlantic and additional body parts were mounted on microscope vent sites include the families Amphilochidae, Stegocepha- slides in Faure’s medium and examined using a Leica MZ lidae, Leucothoidae, Oedicerotidae, Phoxocephalidae, compound microscope with a drawing tube. All dissected Pleustidae, Podoceridae, and Stenothoidae (Bellan-Santini parts were from the left side of each animal. Pencil and Thurston 1996; Bellan-Santini 2005, 2006, 2007; drawings from the microscope were used as the basis for Schander et al. 2010). The amphipod fauna of hydrothermal the line drawings, and were corrected, when necessary, vents normally does not include melitids or maerids, with the using scanning electron microscope (SEM) images. The exception of Bathyceradocus wuzzae found near, but not on, drawings were inked with Adobe Illustrator CS4 following the Wuzza Bare Mount vent field in the NE Pacific (Larsen Coleman (2003, 2009). Setae definitions are based on and Krapp-Schickel 2007). Watling (1989) and Lowry and Stoddart (1992). SEM The amphipod family Melitidae has long been a large images of gills, as well as of the morphology of mouthparts and unwieldy one. Records of Melitidae from abyssal and and legs were produced using a ZEISS Supra 55V SEM on hadal depths are rare (Thurston 2000), and they are also pre-dissected alcohol-dried material, coated with gold/ relatively uncommon in Arctic and Antarctic waters (Jar- palladium. rett and Bousfield 1996; De Broyer et al. 2007). In recent DNA from alcohol-preserved specimens was extracted years, the Maeridae have been separated phylogenetically using the FastDNAÒ SPIN for Soil Kit following the kit from the Melitidae (Krapp-Schickel 2008), and within protocols. A 16SrDNA clone library of gill symbionts was 123 Polar Biol (2012) 35:705–716 707 Fig. 1 a Map with sampling locality.