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A Guide to Extant Coccolithophore Taxonomy Jeremy Young Markus Geisen Lluisa Cros Annelies Kleijne Claudia Sprengel Ian Probert Jette Østergaard

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A Guide to Extant Coccolithophore Taxonomy Jeremy Young Markus Geisen Lluisa Cros Annelies Kleijne Claudia Sprengel Ian Probert Jette Østergaard A guide to extant coccolithophore taxonomy Jeremy Young Markus Geisen Lluisa Cros Annelies Kleijne Claudia Sprengel Ian Probert Jette Østergaard Journal of Nannoplankton Research Special Issue 1 A guide to extant coccolithophore taxonomy Journal of Nannoplankton Research Special Issue 1 ISSN 1210-8049 The Journal of Nannoplankton Research Special Issue is published by the International Nannoplankton Association. Copyright © 2003. The copyright remains with the authors. Printed by Druckstudio Digital Concept, Bremerhaven, Germany. Cover illustration: False coloured image of a Pappomonas sp. from the Alboran Sea, western Mediterranean. Back cover: False coloured image of Coronosphaera mediterranea from the S. Atlantic, off Namibia. A GUIDE TO EXTANT COCCOLITHOPHORE TAXONOMY Jeremy R. Young, Markus Geisen* Palaeontology Dept, The Natural History Museum, Cromwell Road, London, SW7 5BD, UK Lluisa Cros Institut de Ciències del Mar, CSIC, Passeig Marítim 37-49, E-08003 Barcelona, Spain Annelies Kleijne Dept. of Paleoecology and Paleoclimatology, Vrije Universiteit Amsterdam, 1085HV Amsterdam, The Netherlands Claudia Sprengel* FB Geowissenschaften, Universität Bremen, D-28334 Bremen, Germany Ian Probert Laboratoire de Biologie et Biotechnologies Marines, Université de Caen, 14032 Caen, France Jette Buch Østergaard Botanical Institute, University of Copenhagen, Øster Farimagsgade 2D, DK-1353 Copenhagen K, Denmark * currrent address: Alfred Wegener Institute for Polar and Marine Research, Am Handelshafen 12, D-27570 Bremerhaven, Germany Dedication To Peter Westbroek and Jan van Hinte, for inspiring coccolithophorid research, and enriching our lives Journal of Nannoplankton Research Special Issue 1 Foreword Coccolithophores are beautiful organisms and also important ones. They are one of the main groups of marine phytoplankton playing key roles in the marine ecosystem as primary producers and in marine biogeochemistry as producers of organic carbon, carbonate and dimethyl sulphide. In addition they are major sediment formers, key biostratigraphic marker fossils and valuable indicators of palaeoceanographic change. These diverse interests have lead to intensive research on extant coccolithophores over the past decade. Interdisciplinary research has been promoted through the European projects EHUX (Coccolithophorid Dynamics: The European Emiliania huxleyi Programme) and CODENET (Coccolithophorid Evolutionary Biodiversity and Ecology Network) projects. In addition there has been extensive work, especially in Europe and Japan, on coccolithophore communities in the plankton and on fluxes of coccoliths in sediment traps. As a result of this recent research the taxonomy of coccolithophores has advanced significantly over the past decade, i.e. since the seminal syntheses of Jordan & Kleijne (1994) and Jordan et al. (1994). So there is a need for a new synthesis, and especially for an identification guide. There is still work to be done, especially formal description of many now well-established informally described species, but coccolithophores are now one of the most comprehensively described, and most reliably identifiable groups of oceanic microplankton. In consequence they are an ideal group for developing study of the pattern and role of biodiversity in plankton ecology. We hope this identification guide will facilitate such studies, as well as informing palaeontologists studying fossil coccoliths of the nature of the modern biota. Acknowledgements This work is a synthesis of many years work and the authors have been assisted by many colleagues. This has included extended discussion of taxonomy and exchange of samples with other nannoplankton taxonomy enthusiasts, especially Mara Cortes, Doan Nhu Hai, Ric Jordan, Vita Pariente, Christian Samtleben and Alexandra Zeltner. Other colleagues have encouraged the work, assisted with sampling and provided invaluable assistance with electron microscopy, the following list may seem improbably long but all these colleagues have made real contributions for which we are indebted: Laura Arin, Alex Ball, Karl-Heinz Baumann, Chantal Billard, Babette Boeckel, Jörg Bollmann, Paul Bown, Sandra Broerse, Mario Cachão, Daniella Crudeli, Marta Estrada, Claire Findlay, Jacqueline Fresnel, José-Manuel Fortuño, Jan van Hinte, Marc Hockfield, Aude Houdan, Andy Howard, Chris Jones, Michael Knappertsbusch, Ramon Margalef, Linda Medlin, Marie-Helene Noel, Hisatake Okada, Richard Pearce, Patrick Quinn, Alberto Sáez, Blair Steel, Hans Thierstein, Maria Triantaphyllou, Peter Westbroek, Emma Williamson, Patrizia Ziveri. Financial and logistic support has been provided by all of our institutions, recognising the fundamental importance of taxonomy in underpinning all biological research. More specifically the EU TMR CODENET project (Coccolithophorid Evolutionary Biodiversity and Ecology Network) directly funded our collaboration and this volume is a contribution of the project. Image credits The illustrations are primarily the authors but we are very grateful to the following for providing images; Babette Boeckel (Plates 1/6; 6/8, 12; 16/4; 23/3; 25/15; 31/6, 9; 39/10); Marie-Helene Noel (Plate 32/1-3), Vita Pariente (Plates 21/1-3; 23/7, 9, 10, 13; 27/1, 6, 9; 50/1), Tien-Nan Yang (plate36/5-6); Jacqueline Fresnel (Plates 5/10, 9/3-6). J. R. Young et al.: A guide to extant coccolithophore taxonomy Journal of Nannoplankton Research Special Issue 1 A GUIDE TO EXTANT COCCOLITHOPHORE TAXONOMY Table of contents Table of content........................................................................................................................................................................... 1 Introduction................................................................................................................................................................................. 2 Terminology................................................................................................................................................................................ 2 Coccolith crystallography and growth........................................................................................................................................ 2 Species list .................................................................................................................................................................................. 4 Life-cycles................................................................................................................................................................................... 4 Higher classification.................................................................................................................................................................... 6 Conventions used and organisation of the Guide ....................................................................................................................... 6 1. Major heterococcolith groups, except Syracosphaerales........................................................................................................ 8 1.1 Isochrysidales (Noelaerhabdaceae) ................................................................................................................................... 8 1.2 Coccosphaerales - oceanic (Coccolithaceae & Calcidisceae) ......................................................................................... 14 1.3 Coccosphaerales - littoral (Pleurochrysidaceae & Hymenomonadaceae)....................................................................... 22 1.4 Zygodiscales (Helicosphaeraceae & Pontosphaeraceae) ................................................................................................ 26 2. Syracosphaerales (Syracosphaeraceae, Calciosoleniaceae & Rhabdosphaeraceae)............................................................. 30 2.1 Genera with appendages.................................................................................................................................................. 32 2.2 S. nodosa group - BCs muroliths with proximal flange only, XCs flat planoliths .......................................................... 36 2.3 S. pulchra group - BCs with 3 flanges &/or spines, XCs muroliths or domal ................................................................ 42 2.4 S. molischii group - placolith-like BCs ........................................................................................................................... 46 2.5 Syracosphaerales - Genus incertae sedis Coronosphaera............................................................................................... 52 2.6 Calciosoleniaceae ............................................................................................................................................................ 54 2.7 Rhabdosphaeraceae ......................................................................................................................................................... 56 3 Heterococcolith families and genera incertae sedis............................................................................................................... 62 3.1 Alisphaeraceae - Alisphaera and Canistrolithus ............................................................................................................. 62 3.2 Umbellosphaeraceae........................................................................................................................................................ 68
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