Dinoflagellates and Ciliates at Helgoland Roads, North
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Helgol Mar Res (2012) 66:11–23 DOI 10.1007/s10152-010-0242-z ORIGINAL ARTICLE Dinoflagellates and ciliates at Helgoland Roads, North Sea Martin Gu¨nther Joachim Lo¨der • Alexandra Claudia Kraberg • Nicole Aberle • Silvia Peters • Karen Helen Wiltshire Received: 19 July 2010 / Revised: 6 December 2010 / Accepted: 27 December 2010 / Published online: 19 January 2011 Ó Springer-Verlag and AWI 2011 Abstract A monitoring programme for microzooplank- Keywords Dinoflagellates Á Ciliates Á North Sea Á ton was started at the long-term sampling station Monitoring Á Helgoland Roads Á Seasonality ‘‘Kabeltonne’’ at Helgoland Roads (54°11.30N; 7°54.00E) in January 2007 in order to provide more detailed knowl- edge on microzooplankton occurrence, composition and Introduction seasonality patterns at this site and to complement the existing plankton data series. Ciliate and dinoflagellate cell Marine research has a long tradition on Helgoland. Water concentration and carbon biomass were recorded on a temperature has been measured at the Helgoland Roads weekly basis. Heterotrophic dinoflagellates were consid- long-term station ‘‘Kabeltonne’’ (54°11.30N; 7°54.00E) erably more important in terms of biomass than ciliates, since 1873 (Wiltshire and Manly 2004), and biological, especially during the summer months. However, in early chemical and physical parameters have been recorded spring, ciliates were the major group of microzooplankton continuously on a work-daily basis since 1962 (Franke grazers as they responded more quickly to phytoplankton et al. 2004). This makes the Helgoland long-term data food availability. Mixotrophic dinoflagellates played a series one of the longest and most detailed aquatic data secondary role in terms of biomass when compared to sets. Unique to this data set are the phytoplankton species heterotrophic species; nevertheless, they made up an numbers counted work-daily to species level wherever intense late summer bloom in 2007. The photosynthetic possible (Wiltshire and Du¨rselen 2004). Since 1975, the ciliate Myrionecta rubra bloomed at the end of the sam- time series also includes meso- and macrozooplankton pling period. Due to its high biomass when compared to determined to species level three times per week (Greve crustacean plankton especially during the spring bloom, et al. 2004). Thus, the time series provides an excellent microzooplankton should be regarded as the more impor- basis for analyses of long-term trends including changes tant phytoplankton grazer group at Helgoland Roads. evinced in the North Sea pelagic system over the recent Based on these results, analyses of biotic and abiotic fac- decades (Schlu¨ter et al. 2008; Wiltshire et al. 2008). Fur- tors driving microzooplankton composition and abundance thermore, it is a very important basis for the parameteri- are necessary for a full understanding of this important sation and validation of mathematical ecosystem models component of the plankton. and is invaluable in biodiversity and global change con- siderations (Wirtz and Wiltshire 2005). However, one important group of planktonic organisms is under-repre- Communicated by H.-D. Franke. sented in the long-term series so far—the microzooplank- ton. Although data on heterotrophic dinoflagellates exist in M. G. J. Lo¨der (&) Á A. C. Kraberg Á N. Aberle Á S. Peters Á the data set, they were recorded with varying degrees of K. H. Wiltshire accuracy (Wiltshire and Du¨rselen 2004) and did not always Biologische Anstalt Helgoland, Alfred Wegener Institute mirror the diversity in species composition (Hoppenrath for Polar and Marine Research, Marine Station, POB 180, 27483 Helgoland, Germany 2004). Long-term data on ciliates, another crucial micro- e-mail: [email protected] zooplankton group, are totally lacking at Helgoland Roads. 123 12 Helgol Mar Res (2012) 66:11–23 The term microzooplankton refers to the size fraction programme (Myrinecta rubra 1999, Laboea strobila 2007, of heterotrophic planktonic organisms between 20 and Mesodinium pulex 2008), no ciliate species had continu- 200 lm. Consisting of a diverse array of protozoa and ously been recorded previously at Helgoland. Therefore, metazoa, its numerically most important components are the new microzooplankton monitoring presented here was heterotrophic dinoflagellates and ciliates (Capriulo et al. to provide completely new information on ciliate biomass 1991). Recent research, for example, by Landry and Calbet and seasonality patterns at a hitherto unavailable taxo- (2004), demonstrated the fundamental importance of nomic resolution. microzooplankton as phytoplankton grazers. They showed The loss of microzooplankton species due to fixative that grazing by microzooplankton can be as high as problems has often been discussed in the literature 60–75% of the daily phytoplankton production. Further- (Stoecker et al. 1994). Thus, we diverged from the neutral more, results indicate that microzooplankton tends to sur- fixative used for the long-term monitoring and used acidic pass mesozooplankton as primary consumers (Sherr and Lugol’s solution (final concentration 2%, Throndsen 1978), Sherr 2007). A meta-analysis of Calbet and Landry (2004) as this is the standard fixative used in most studies on revealed that microzooplankton grazing can be responsible microzooplankton composition. The concentration we used for 60% of phytoplankton mortality in coastal and estuarine has been proven to be the best compromise for both con- environments (comparable to Helgoland Roads in terms of serving higher concentrations of ciliates and preventing chlorophyll a concentration). We realised from observa- shrinkage of cells (Stoecker et al. 1994). A subsample of tions that microzooplankton could also potentially be the 250 mL was fixed immediately. Samples were stored in the most important grazer group in waters at Helgoland. cold and dark, and then 50 mL of the sample were settled Therefore, the aim of this study was to supplement the for 24 h and counted under an inverted microscope (Zeiss regular plankton monitoring series at Helgoland with an Axiovert 135) using the Utermo¨hl method (Utermo¨hl 1958, intensive monitoring of the microzooplankton. Investiga- Lund et al. 1958). At least half of the surface of the sedi- tions into species composition and seasonality of this mentation chamber or the whole chamber was counted out important functional grazer group (dinoflagellates and cil- at 200-fold magnification, thus reducing counting biases iates) on a more regular basis will provide vital baseline against rare species. Identification of naked dinoflagellates data for studies of long-term changes in the microzoo- and especially of ciliates in Lugol’s solution-preserved plankton community and the pelagic system at Helgoland. samples is often difficult below genus level (Johansson et al. 2004), even with the modified fixation method applied here. Therefore, problematic ciliates and dino- Materials and methods flagellates were identified to genus level or, otherwise, pooled into size-dependent groups and ‘‘morphotypes’’, A 2.5-year microzooplankton monitoring programme has based on their similar shape. Mixotrophy of the ciliates was been carried out at Helgoland Roads to investigate the not measured; thus, we have no exact data on the per- abundance of dinoflagellates and ciliates in the Southern centage of mixotrophic ciliates in the samples. However, to North Sea. This monitoring hoped to establish a higher date, all mixotrophic ciliates have been shown to be taxonomic resolution and to improve the evaluation of phagotrophic (Sherr and Sherr 2002) and consequently all biomass for single taxa of microzooplankton. ciliates except Myrionecta rubra could be considered het- From January 2007 until June 2009, samples were taken erotrophic (Johansson et al. 2004). The latter species acts once a week at the ‘‘Kabeltonne’’ site (54°11.30N; 7°54.00E) essentially as a phototroph (Montagnes et al. 2008), but as at Helgoland. These data supplemented the routine sam- recent studies have shown that it also has phagotrophic pling programme, which is carried out week-daily and for capabilities (Park et al. 2007), we thus recorded it as which plankton samples are fixed with a weak neutral mixotrophic. Lugol’s solution (final concentration 0.5%) (Wiltshire et al. As most chloroplast-bearing dinoflagellates are also 2008). Although dinoflagellates are counted within the capable of mixotrophic nutrition via phagotrophy (Du Yoo long-term programme, the taxonomic focus lies on phyto- et al. 2009), these were considered to be potential grazers plankton groups such as diatoms. Due to the time-con- with more or less marked phagotrophic capabilities. The suming counting procedure for phytoplankton and the high identification of dinoflagellates was primarily based on frequency of samples (work-daily), rare, small or uncom- Dodge (1982), Tomas (1996) and Hoppenrath et al. (2009). mon dinoflagellate taxa are inevitably neglected or cate- Ciliates were determined based on Kahl (1932), Carey gorised into size classes. The present study investigated (1992) and Montagnes (2003). such under-represented species more intensely during the As an additional new feature compared to the regular long- 2.5-year microzooplankton monitoring. Apart from three term series, each taxon recorded during counting was docu- ciliates that have recently been included in the counting mented by an image (archived online at http://planktonnet. 123 Helgol Mar Res (2012) 66:11–23 13 awi.de). These images, used for subsequent biovolume esti- between 0.2