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Zooplankton of the Open Baltic Sea: Extended Atlas LEIBNIZ-INSTITUT FÜR OSTSEEFORSCHUNG WARNEMÜNDE LEIBNIZ INSTITUTE FOR BALTIC SEA RESEARCH Meereswissenschaftliche Marine Science Berichte Reports No. 76 2009 Zooplankton of the Open Baltic Sea: Extended Atlas by Irena Telesh, Lutz Postel, Reinhard Heerkloss, Ekaterina Mironova, Sergey Skarlato "Meereswissenschaftliche Berichte" veröffentlichen Monographien und Ergebnis- berichte von Mitarbeitern des Leibniz-Instituts für Ostseeforschung Warnemünde und ihren Kooperationspartnern. Die Hefte erscheinen in unregelmäßiger Folge und in fortlaufender Nummerierung. Für den Inhalt sind allein die Autoren verantwortlich. "Marine Science Reports" publishes monographs and data reports written by scien- tists of the Leibniz-Institute for Baltic Sea Research Warnemünde and their co- workers. Volumes are published at irregular intervals and numbered consecutively. The content is entirely in the responsibility of the authors. Schriftleitung: Dr. Lutz Postel ([email protected]) Bezugsadresse / address for orders: Leibniz–Institut für Ostseeforschung Warnemünde Bibliothek Seestr. 15 D-18119 Warnemünde Germany ([email protected]) Eine elektronische Version ist verfügbar unter / An electronic version is available on: http://www.io-warnemuende.de/research/mebe.html This volume is listed as The Baltic Marine Biologists Publication No.21 and should be cited: Telesh, I., Postel, L., Heerkloss, R., Mironova, E., Skarlato, S., 2009. Zooplankton of the Open Baltic Sea: Extended Atlas. BMB Publication No. 21 – Meereswiss. Ber., Warnemünde, 76, 1 - 290. ISSN 0939 -396X Meereswissenschaftliche Berichte MARINE SCIENCE REPORTS No. 76 Zooplankton of the Open Baltic Sea: Extended Atlas By Irena Telesh1, Lutz Postel2, Reinhard Heerkloss3 Ekaterina Mironova4, Sergey Skarlato4 1 Zoological Institute, Russian Academy of Sciences, Universitetskaya Emb. 1, 199034 St. Petersburg, Russia 2 Leibniz – Institute for Baltic Sea Research (IOW), Seestr. 15, D-18119 Rostock-Warnemünde, Germany 3 Institute for Biological Science, University of Rostock, Albert-Einstein-Str. 3, D-18051 Rostock, Germany 4 Institute of Cytology, Russian Academy of Sciences, Tikhoretsky Ave. 4, 194064 St. Petersburg, Russia Corresponding author: [email protected] Leibniz – Institut für Ostseeforschung Warnemünde 2009 In memory of Professor Ulrich Schiewer CONTENTS Page Preface…………………………………...……………….…….…………7 1. Introduction………………………………………………..…………..9 2. General characteristics of zooplankton of the Baltic Sea……….........15 3. Methods of collecting and analysing zooplankton in the Baltic Sea………...……………………………………….............................21 3.1. Sampling: general aspects….………………………………...21 3.2. Sampling of meso- and macrozooplankton…………………..23 3.3. Sampling and study of microzooplankton................................27 3.4. Identification and counting of meso- and macrozooplankton……………………………………………..28 3.5. Biomass determination.............................................................32 3.6. Picture key to major zooplankton taxa…….…………….…...33 4. Ciliates of the Baltic Sea……………………………………...…...…35 4.1. Brief characteristics of planktonic ciliates of the Baltic Sea………………………………………………………………...35 4.2. Checklist of ciliates of the Baltic Sea……………………......43 4.3. Photo plates: ciliates of the Baltic Sea….…..……….…….....81 5. Meso- and macrozooplankton of the open Baltic Sea...………...…..123 5.1. Description of most abundant meso- and macrozooplankton groups……………………………………….123 5.2. Checklist of meso- and macrozooplankton of the open Baltic Sea……………………………………………………..….149 5.3. Photo plates: meso- and macrozooplankton of the Baltic Sea………………………………………………..……………....161 Acknowledgements…..………………..………………………………..249 References....……………………………….………………………..….251 List of selected zooplankton Internet data bases………….…………….267 Index of Latin names…………………………………….…………...…269 5 ABSTRACT This is the Second Edition (improved and extended) of the third volume in a series of zooplankton atlases of the Baltic Sea. It describes zooplankton community of the open Baltic waters, which is a mixture of marine species and diverse representatives of brackish water and limnetic faunas. Brief information on morphology and ecology of zooplankters, picture key to higher invertebrate taxa and methodological recommendations for sampling, identification and counting of zooplankton are provided. Two checklists present more than 1030 names of micro-, meso- and macrozooplankton organisms. The volume is enhanced by 4 tables, 31 line drawing, and 462 colour photographs. The photographs of zooplankters are combined into 63 photo plates depicting most common species of Protozoa (Ciliata), Cnidaria, Ctenophora, Turbellaria, Rotifera, Phyllopoda, Copepoda, Chaetognatha and Copelata, as well as meroplanktonic larvae of Polychaeta, Mollusca, Cirripedia, Bryozoa and Echinodermata. The atlas is recommended for university students, zoologists and aquatic ecologists who are investigating and monitoring the pelagic ecosystem of the Baltic Sea. 6 PREFACE This volume is the Second Edition (revised and extended by new information on ciliates) of the third book in a series of zooplankton atlases – an illustrated inventory of planktonic invertebrates inhabiting the open waters of the Baltic Sea. The two previous volumes deal with the estuarine mesozooplankton of the Baltic Sea (Part I, Rotifera: Telesh & Heerkloss, 2002; Part II, Crustacea: Telesh & Heerkloss, 2004). The necessity of creating a zooplankton inventory of the Baltic Sea had been under discussion since the 1980-s when Dr. Gerda Behrends headed the Working Group (WG) “Zooplankton” within the association of the Baltic Marine Biologists (BMB). In the early 2000-s, the BMB WG No. 29 “Zooplankton Diversity” was established (convener – Dr. Irena Telesh) in order to continue and facilitate the biodiversity research in the Baltic Sea region. The present series of zooplankton atlases of the Baltic Sea was initiated and produced by the members of the BMB WG No. 29 “Zooplankton Diversity”. It is worth mentioning here that since 1998 Professor Ulrich Schiewer (1936-2007) from the University of Rostock, to whom we dedicate the present volume, enthusiastically supported this work. This atlas provides information on the zooplankton of the open Baltic Sea, which is to great extent a mixture of marine species and diverse brackish water and limnetic faunas typical for the vast estuarine and coastal areas located mainly in the southern and north-eastern parts of the Baltic. The specific feature of this volume is the updated species list of meso- and macrozooplankton of the open Baltic (217 species), and the checklist of ciliates (814 species) inhabiting open waters of both, central and coastal areas of the Baltic Sea. The atlas is illustrated by 31 line drawing and 462 colour photographs. The photographs are combined into 63 photo plates depicting most common holo- and meroplanktonic representatives of different invertebrate taxa: Protozoa (Ciliata), Cnidaria, Ctenophora, Turbellaria, Rotifera, Phyllopoda, Copepoda, Cirripedia, Polychaeta, Mollusca, Bryozoa, Echinodermata, Chaetognatha and Copelata. A new feature of the present edition is the extended chapter “Ciliates of the Baltic Sea” which contributes substantially to both biodiversity and cyto- ecological research directions in the area. The chapter is illustrated by 197 original colour photographs that describe the diverse cell morphology of the free-living ciliated protists in the Baltic Sea. Photographs of live ciliates were made with the help of the Research Microscope LEICA DM-2500 at magnifications 40x and 100x using the 7 regimes of bright field (BF), differential interference contrast (DIC), or phase- contrast (PHC). Original photographs of major other zooplankters were made with a CANON video system connected to OLYMPUS Research Microscope BH-2. In the illustrations, the magnification is indicated by a scalebar with a number which is its length in micrometers (μm). Alternatively, the size of a depicted organism is indicated in the legend. This volume should be of practical use for students and technicians as well as for scientists. Suggestions about how to improve the project and cooperation to enhance it are most welcome. 8 1. INTRODUCTION Since the last glaciation, the Baltic Sea has undergone several evolutionary stages between a huge marine bay and a large freshwater lake during which a number of different ecological systems developed and were successfully replaced in this area (Jansson, 1972). Today the Baltic Sea is the largest brackish water area in the world characterized as a temperate shelf sea with permanent salinity stratification, a horizontal salinity gradient and low water turnover of 35 years. It is a shallow sea with a mean depth of 62.1 m, the greatest depth 459 m, an area of 415,266 km2 (Baltic Proper itself is 211,069 km2), a volume of ca. 22,000 km3 (HELCOM, 2001; Wulff et al., 2001; Schiewer, 2008). The presence of shallow sills at the western inlets causes stable water stratification. The Baltic can be best compared to a stratified fiord with a rich supply of fresh water from the rivers. Due to the humid climate, there is an estuarine circulation with outflow of low-saline water above the halocline and powerful periodic injections of North Sea water below the halocline which greatly affect the salinity of the deep water layers. The location of the Baltic Sea in the northern high latitudes means a pronounced seasonality in temperature and light regime. The vegetation season lasts longer in the southern
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