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Zooplankton of the open Baltic: Extended Atlas - IOW

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4. CILIATES OF THE BALTIC SEA<br />

4.1. Brief characteristics <strong>of</strong> planktonic ciliates <strong>of</strong> <strong>the</strong> <strong>Baltic</strong> Sea<br />

General information<br />

Planktonic ciliates play pivotal roles in <strong>the</strong> transfer and recycling <strong>of</strong><br />

carbon and o<strong>the</strong>r nutrients in <strong>the</strong> seas. It is now well established that ciliates<br />

constitute an essential food link in marine environments since <strong>the</strong>y remove a<br />

significant part <strong>of</strong> primary and bacterial production and assume ‘trophic<br />

repackaging’ <strong>of</strong> picoplankton particles (0.2–2.0 µm) o<strong>the</strong>rwise inaccessible to<br />

mesozooplankton. Picoplankton organisms (heterotrophic bacteria,<br />

cyanobacteria, phytoplankton <strong>of</strong> cell size 0.2–2.0 µm) are <strong>the</strong> major<br />

consumers <strong>of</strong> dissolved organic matter; <strong>the</strong>ir biomass and production is <strong>the</strong><br />

largest within <strong>the</strong> total planktonic community. Ciliates, which are among <strong>the</strong><br />

most important consumers <strong>of</strong> this production, play a key role as an<br />

intermediate stage in <strong>the</strong> transformation <strong>of</strong> organic matter from picoplankton<br />

to higher trophic levels.<br />

Firstly, ciliates have high growth and metabolic rates. Secondly, <strong>the</strong>y<br />

belong to a size range easily removed by metazoans, <strong>the</strong>refore <strong>the</strong>y constitute<br />

a significant part <strong>of</strong> <strong>the</strong> food biomass in <strong>the</strong> sea. Planktonic ciliates make an<br />

essential contribution to <strong>the</strong> nutrition <strong>of</strong> copepods, rotifers, o<strong>the</strong>r invertebrates<br />

and fish larvae. Along with phototrophic microorganisms, viruses,<br />

heterotrophic bacteria, and microscopic fungi, ciliates participate in a<br />

‘microbial loop’, which is an integral part <strong>of</strong> <strong>the</strong> planktonic food webs.<br />

Fur<strong>the</strong>rmore, <strong>the</strong> importance <strong>of</strong> planktonic ciliates as indicators <strong>of</strong><br />

eutrophication and water pollution has been emphasized.<br />

History <strong>of</strong> research<br />

Pioneering studies <strong>of</strong> planktonic ciliates in <strong>the</strong> <strong>Baltic</strong> Sea began in <strong>the</strong><br />

late 1940-s (Biernacka, 1948). Most <strong>of</strong> <strong>the</strong>m dealt with Tintinnids because <strong>of</strong><br />

<strong>the</strong>ir relatively large size (Biernacka, 1948, 1952; Hedin, 1974, 1975). There<br />

were also researches <strong>of</strong> o<strong>the</strong>r groups <strong>of</strong> ciliates (Bock, 1960; Biernacka,<br />

1963). Investigations <strong>of</strong> benthic ciliates began earlier, in <strong>the</strong> late 1920-s<br />

(Sauerbrey, 1928; Kahl, 1930-1935). Contemporary investigations <strong>of</strong> <strong>Baltic</strong><br />

ciliates deal with both, benthic communities (Klinkenberg & Schumann,<br />

1994; Dietrich & Arndt, 2000; Garstecki et al., 2000), and planktonic ciliates<br />

(Smetacek, 1981; Boikova, 1989; Arndt, 1991; Kivi & Setala, 1995; Uitto et<br />

al., 1997; Witek, 1998; Setala & Kivi, 2003; Johansson et al., 2004;<br />

Samuelsson et al., 2006; Beusekom et al., 2007). Benthic and pelagic<br />

communities <strong>of</strong> ciliates showed little taxonomic overlap (Garstecki et al.,<br />

2000). However, benthic ciliates (genus Euplotes, Aspidisca, Tracheloraphis<br />

etc.) may be found in <strong>the</strong> water column because <strong>of</strong> bottom hashing; <strong>the</strong>ir role<br />

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