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Taxonomy and Ecology of the Ciliate Fauna (Protozoa, Ciliophora) in the Endopagial and Pelagial of the Weddell Sea, Antarctica

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Taxonomy and Ecology of the Ciliate Fauna (Protozoa, Ciliophora) in the Endopagial and Pelagial of the Weddell Sea, Antarctica © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at Taxonomy and ecology of the ciliate fauna (Protozoa, Ciliophora) in the endopagial and pelagial of the Weddell Sea, Antarctica Wolfgang PETZ, Weibo SONG & Norbert WILBERT Stapfia 40 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at Stapfia 40 223 pp. 23.8.1995 Taxonomy and ecology of the ciliate fauna (Protozoa, Ciliophora) in the endopagial and pelagial of the Weddell Sea, Antarctica 3 Wolfgang PETZ1'2, Weibo SONG & Norbert WILBERT1 1 Institut fur Zoologie, Universität Bonn, Poppelsdorfer Schloss, D-53115 Bonn, Germany. 2 Institut für Zoologie, Universität Salzburg, Hellbrunner Strasse 34, A-5020 Salzburg, Austria. 3 College of Fisheries, Ocean University of Qingdao, Qingdao 266003, P. R. China. Correspondence to: Dr. Wolfgang PETZ Universität Salzburg, Institut für Zoologie, Hellbrunner Strasse 34, A-5020 Salzburg, Austria email: [email protected] © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at Contents Introduction 4 Material and methods 4 Type material 7 Results and discussion 7 Colonization of sea ice 7 Methodological considerations 10 Terminology 10 Faunistics 11 Description of species 12 Order Prostomatida SCHEWIAKOFF, 1896 15 Genus Pseudotrachelocerca SONG, 1990 15 Genus Placus COHN, 1866 18 Order Spathidiida FOISSNER & FOISSNER, 1988 21 Genus Didinium STEIN, 1859b 21 Genus Chaenea QUENNERSTEDT, 1867 24 Genus Fuscheria FOISSNER, 1983 29 Genus Lacrymaria BORY DE ST. VINCENT, 1826 32 Order Cyclotrichida JANKOWSKI, 1980 40 Genus Myrionecta JANKOWSK1, 1976 40 Genus Rhabdoaskenasia KRMNER & FOISSNER, 1990 41 Order Pleurostomatida SCHEWIAKOFF, 1896 43 Genus Litonotus WRZESNIOWSKI, 1870 43 Genus Kentrophyllum nov. gen 50 Genus Loxophyllum DUJARDIN, 1841 55 Order Synhymeniida PUYTORAC et al., 1974b 59 Genus Zosterodasys DEROUX, 1978 59 Order Cyrtophorida FAURE-FREMIET in CORLISS, 1956 62 Genus Chlamydonella nov. gen 62 Genus Dysteria HUXLEY, 1857 66 Family Kryoprorodontidae ALEKPEROV & MAMAJEVA, 1992 68 Genus Gymnozoum MEUNIER, 1910 69 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at Order Hymenostomatida DELAGE & HEROUARD, 1896 79 Genus Frontonia EHRENBERG, 1838 79 Order Scuticociliatida SMALL, 1967 83 Genus Cryptochilum MAUPAS, 1883 83 Genus Uronema DUJARDIN, 1841 88 Genus Pleuronema DUJARDIN, 1836 97 Genus Porpostoma MOEBIUS, 1888 101 Order Heterotrichida STEIN, 1859a 103 Genus Condylostoma BORY DE ST. VINCENT, 1826 103 OrderStrombidiidaJANKOWSKI, 1980 Ill GenusStrombidiumCLAPABEDE& LACHMANN, 1859 Ill Genus Spirostrombidium JANKOWSKI, 1978 123 Genus Tontonia FAURE-FREMIET, 1914 130 Order Oligotrichida BUETSCHLI, 1887 134 Genus Leegaardiella LYNN &MONTAGNES, 1988 134 Genus Pelagostrobilidiumnov. gen 138 Genus Rimostrombidium JANKOWSKI, 1978 141 Genus Codonellopsis JOERGENSEN, 1924 144 Genus Laackmanniella KOFOID& CAMPBELL, 1929 151 Genus Cymatocylis LAACKMANN, 1910 154 Order Hypotrichida STEIN, 1859a 159 Genus Holosticha WRZESNIOWSK1, 1877 159 Genus Notocephalus nov. gen 169 Genus Aspidisca EHRENBERG, 1830 172 Genus Uronychia STEIN, 1859c 173 Genus Cytharoides TUFFRAU, 1974 178 Genus Diophrys DUJARDIN, 1841 182 Genus Euplotes EHRENBERG, 1830 183 Other ciliates 197 Summary 198 Acknowledgements 199 References 200 Systematic index 219 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at Introduction Every winter, the ice sheet around Antarctica increases from about 4xlO6 km2 up to 6 2 20x10 km by freezing of the Southern Ocean (MAYKUT 1985). Sea ice is, however, not a compact structure but interlaced with brine-filled pores and channels about 200 urn to some cm in diameter (WEISSENBERGER et al. 1992). This internal system is colonized by an abundant community of bacteria, fungi, algae (mainly diatoms), protozoans and small metazoans (Fig. 2; e.g. GARRISON 1991; GARRISON & BUCK 1989b, 1991; GROSSMANN & DIECKMANN 1994; HORNER 1985; KOTTMEIER & SULLIVAN 1987; PALMISANO & GARRISON 1993; PALMISANO & SULLIVAN 1983; SPINDLER 1994; SPINDLER & DIECKMANN 1991; SPINDLER et al. 1990; STOECKER et al. 1990, 1993). Already early Antarctic pioneers noticed the distinctly brownish coloured layer in sea ice which marks the intensively populated zone (HOOKER 1847). A more detailed biological survey started only recently and revealed that the ice biota is of global significance (LEGENDRE et al. 1992). With very few exceptions (AGATHA et al. 1990, 1993; CORLISS & SNYDER 1986; FENCHEL & LEE 1972; PETZ 1994, 1995; WILBERT et al. 1993), the composition of the ice ciliate fauna has not been investigated before and is thus only very incompletely known. A comprehensive taxonomic study of sea ice and planktonic ciliates was thus performed in situ utilizing detailed in vivo observations, hi addition, the colonization of sea ice was investigated in the austral autumn. To provide a sound basis for the identification of Antarctic sea ice ciliates by non-specialists, all the species found are described and illustrated in detail. Material and methods Sea ice samples were collected using an ice-coring auger of 7.5 cm or 10 cm diameter, then cut in 10-cm-sections, melted overnight at +1°C in about the same volume of sterile filtered seawater to relieve osmotic stress for the organisms and subsequently immediately investigated using a cooled microscope. Grease ice and very young pancake ice was obtained using an ice basket. For details on ice formation and types see, e.g., LANGE et al. (1989), MAYKUT (1985). © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at -90° 0 C E A N 0 500 1000 Fig. 1: Study area (arrowhead). Active ciliate numbers were estimated using a direct live counting method (detailed description in LÜFTENEGGER et al. 1988): 8 subsamples ä 0.25 ml were examined from each melted sample; cihate abundances are already corrected for dilution. Plankton catches were made with an Apstein-net (mesh size 20 um) from 0-20 m depth for qualitative, and with a bottle sampler (bio-rosette) from 0, 10 and 50 m for quantitative studies (4 x 0.25 ml per sample). The MANN-WHITNEY U-test was computed following KÖHLER et al. (1984). Field material and raw cultures were used for morphologic studies. WiLBERT's (1975) protargol silver impregnation technique was applied to reveal the infraciliature, the CHATTON-LWOFF silver nitrate method according to CORLISS (1953) was used to stain the silverline system. Preparation for scanning electron microscopy follows the FoiSSNER (1991) protocol. Living individuals were measured at X 100 magnification; counts and measurements on stained specimens were performed at X 400 and X 1000 magnification (1 measuring unit = 3.1 urn and 1.2 urn, respectively). All © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at morphometric data are based on randomly selected, protargol impregnated and mounted non-dividers; statistics were calculated following SOKAL & ROHLF (1981); abbreviations in the tables: CV, coefficient of variation in %; M, median; Max, maximum; Min, minimum; SD, standard deviation; SE, standard error of arithmetic mean; x, arithmetic mean. Morphologic terminology follows mainly BORROR (1972a), CORLISS (1979), DEROUX (1970, 1976b), FOISSNER (1984a), KAHL (1930, 1931, 1932) and WALLENGREN (1900) and the systematic classification CORLISS (1979). Impregnated cells were drawn using a camera lucida. Unless otherwise noted, biomass was estimated from biovolume using in vivo dimensions, i.e. 1 urn3 = 1 pg protoplasm (FiNLAY 1982), and simple geometric bodies. The investigation was performed during the cruise ANT X/3 of the RV „Polarstern" in the eastern Weddell Sea in April and May 1992 (Fig. 1). Fig. 2: Sea ice biotopes (pagial). En, endopagial; Ep, epipagial; Hy, hyperpagial; Me, metapagial; Pc, pancake; Pg, pagiotelma; Pi, drifting pack ice. © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at Type material As noted, holotype, paratype and neotype slides as well as preparations of other species for reference have been deposited in the collection of microscope slides of the Oberösterreichische Landesmuseum (LI), A-4040 Linz, Austria. Neotypes have been designated for Codonellopsis glacialis, Condylostoma grcmulosum, Cymatocylis calyciformis, C. convallaria, Didinium gargantua, Lacrymaria lagenula, Loxophyllum rostratum, Strombidium antarcticum, S. crassulum, S. emergens and Uronychia transfuga. In the course of this study, types of Euplotes algivorus and original slides of Cohnilembus grassei, Spiroprorodon garrisoni, Strombidium rhyticollare and Tachysoma parvulum were investigated; types of the latter species have not yet been deposited. The type slides of Spiroprorodon glacialis are apparently lost (see below). Results and discussion Colonization of sea ice Only few active ciliates were found in the water column with the quantitative sampling procedure (Table 1). Grease ice (initial stage of freezing) and very young pancake ice (next stage of sea ice formation) also contained no or very few active ciliates, usually planktonic species (Tables 1,2). Considerably higher numbers (up to 31 173 active ind./l melted ice) were found in slightly older (max. 50 days; GRADINGER et al. 1993), about 40-cm-thick pancake ice (Table 1). This suggests that a rapid colonization or fast population growth of ciliates occurs within the ice. Growth in sea ice samples is indicated by the regular observation of dividers of, e.g., strombidiids, thigmokeronopsids, Aspidisca
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