Scrippsiella Trochoidea (F.Stein) A.R.Loebl
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MOLECULAR DIVERSITY AND PHYLOGENY OF THE CALCAREOUS DINOPHYTES (THORACOSPHAERACEAE, PERIDINIALES) Dissertation zur Erlangung des Doktorgrades der Naturwissenschaften (Dr. rer. nat.) der Fakultät für Biologie der Ludwig-Maximilians-Universität München zur Begutachtung vorgelegt von Sylvia Söhner München, im Februar 2013 Erster Gutachter: PD Dr. Marc Gottschling Zweiter Gutachter: Prof. Dr. Susanne Renner Tag der mündlichen Prüfung: 06. 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München, 28.02.2013 ……..……………………….......... 4 5 ACKNOWLEDGEMENTS I would like to express my gratitude to Prof. Dr. Helmut Keupp and my doctoral thesis supervisor PD Dr. Marc Gottschling for the enabling to conduct my PhD thesis on the highly interesting topic of marine Dinophytes. The project was funded by the DFG grant Ke 322/36. Prof. Keupp kindly helped with words and deeds to negotiate administrative barriers. Marc was always a dedicated and patient mentor, who supported me anytime in difficulties and challenges. I also would like to thank Prof. Dr. Susanne Renner for hosting me at the LMU Munich during the last four years, for many inspiring lab group meetings and for kindly being a referee of my PhD thesis. I am grateful to Prof. Dr. Jörg Nickelsen, Prof. Dr. Herwig Stibor, Prof. Dr. Günther Heubl and PD Dr. Martin Heß, who kindly read and marked my PhD thesis. I appreciated the collaborations and cooperation with Dr. Eva Facher (LMU München), Dr. Haifeng Gu (Third Institute of Oceanography, Xiamen, China), Dr. Uwe John (AWI, Bremerhaven), Monika Kirsch (Paleontology, University of Bremen), Prof. Dr. Kazumi Matsuoka (Nagasaki University, Japan), Dr. K.J. Sebastian Meier (Paleontology, University of Kiel), Prof. Dr. Alexandros Stamatakis (KIT, Karlsruhe), Dr. Urban Tillmann (AWI, Bremerhaven), Dr. Thomas von Rintelen (Museum für Naturkunde Berlin) and Dr. Carmen Zinßmeister (Senckenberg am Meer, Wilhelmshaven). My special thanks goes to Agnes, Andi, Chen, Flo, Juan Carlos, my family and friends, who supported me greatly in the last stressful phase of writing and defending my thesis. 6 PUBLICATIONS & MANUSCRIPTS PAPER 1 (PUBLICATION) Zinssmeister, C., Soehner, S., Kirsch, M., FAcher, E., Meier, K.J.S., Keupp, H. & Gottschling, M. (2012) SAme but different: Two novel bicArinAte species of extAnt cAlcAreous dinophytes (ThorAcosphAerAceAe, PeridiniAles) from the MediterrAneAn SeA. J. Phycol. 48 (5): 1107-1118. Own contributions: Fieldwork (40%); moleculAr lAb work (100%); moleculAr phylogenetic anAlysis (incl. imAges 80%); totAl mAnuscript prepArAtion (20%). PAPER 2 (MANUSCRIPT) Gu, H., Kirsch, M., Zinssmeister, C., Soehner, S., Meier, K.J.S., Liu, T. & Gottschling, M. WAking the deAd: MorphologicAl And moleculAr chArActerizAtion of extAnt Posoniella tricarinelloides (ThorAcosphAerAceAe, DinophyceAe). Protist (in press). Own contributions: Fieldwork (20%); moleculAr lAb work (50%); moleculAr phylogenetic AnAlysis (incl. imAges 10%); totAl mAnuscript prepArAtion (10%). PAPER 3 (PUBLICATION) TillmAnn, U., Soehner, S., NézAn, E. & Krock, B. (2012) First record of the genus Azadinium (DinophyceAe) from the ShetlAnd IslAnds, including the description of Azadinium polongum sp. nov. HArmful AlgAe 20: 142-155. Own contributions: MoleculAr lAb work (50%); moleculAr phylogenetic AnAlysis (incl. imAges 100%); totAl mAnuscript prepArAtion (20%). 7 PAPER 4 (PUBLICATION) Zinssmeister, C., Soehner, S., Facher, E., Kirsch, M., Meier, K.J.S. & Gottschling, M. (2011) Catch me if you can: The taxonomic identity of Scrippsiella trochoidea (F.Stein) A.R.Loebl. (Thoracosphaeraceae, Dinophyceae). Syst. Biodivers. 9: 145-157. Own contributions: Fieldwork (40%); molecular lab work (100%); molecular phylogenetic analysis (incl. images 60%); total manuscript preparation (20%). PAPER 5 (PUBLICATION) Soehner, S., Zinssmeister, C., Kirsch, M. & Gottschling, M. (2012) Who am I – and if so, how many? Species diversity of calcareous dinophytes (Thoracosphaeraceae, Peridiniales) in the Mediterranean Sea. Org. Divers. Evol. 12 (4): 339-348. Own contributions: Fieldwork (40%); molecular lab work (100%); network analysis (incl. images 100%); total manuscript preparation (70%). PAPER 6 (PUBLICATION) Gottschling, M., Soehner, S., Zinssmeister, C., John, U., Plötner, J., Schweikert, M., Aligizaki, K. & Elbrächter, M. (2012) Delimitation of the Thoracosphaeraceae (Dinophyceae), including the calcareous dinoflagellates, based on large amounts of ribosomal RNA sequence data. Protist 163: 15-24. Own contributions: Molecular lab work (50%); molecular phylogenetic analysis (incl. images 30%); total manuscript preparation (20%). 8 9 GENERAL INTRODUCTION THE STUDY GROUP – THE DINOPHYCEAE The Dinophyceae (‘Dinoflagellata’ under the zoological code of nomenclature: ICZN) are an ecologically important and evolutionarily fascinating group of protists. The majority of Dinophyceae, comprising approximately 2000 described extant species, are free-living, marine organisms, occuring from polar through temperate to tropical waters (Taylor et al. 2008; Gómez 2012). These unicellular algae are ancient organisms with fossil record dated back into the Silurian, over 400 million years ago (Steidinger & Tangen 1996). In the second half of the 18th century, the first extant dinophytes have been described (Moestrup & Daugbjerg 2007). From the end of the 1950s onwards, the Dinophyceae increasingly attracted the researchers’ interest (Figure 1), which emphasises the ecological and economical importance of this group. Together with the Apicomplexa and the Ciliata, the Dinophyceae belong to the Alveolata, which are characterised by specialised vesicles located directly beneath the cell surface and called alveoli (Cavalier- Smith 1993; Fast et al. 2002; Leander & Keeling 2004). Morphological Figure 1: Number of publications per year (Web of Knowledge data base search on “dinofl*” and “dinophy*”). apomorphies of the dinophytes are the unique dinokaryon with permanently condensed, liquid-crystalline chromosomes and the lack of typical eukaryotic histones or nucleosomes (Spector 1984; Moreno Díaz de la Espina et al. 2005; Lin 2011). The biflagellate cells possess a coiled transversal flagellum positioned in an encircling groove coined “cingulum” (allowing locomotion) and a longitudinal flagellum (giving propulsive force) (Taylor 1980; Hackett et al. 2004). The monophyly of the Dinophyceae is highly supported based on several molecular loci (Maroteaux et al. 1985; Saldarriaga et al. 2003; Shalchian- Tabrizi et al. 2006a; Zhang et al. 2007; Minge et al. 2009; Hoppenrath & Leander 2010). 10 Figure 2: The Dinophyceae are highly diverse in morphology. This simplified phylogeny indicates the classification based on the arrangement of thecal plates: dinophysioid, gonyaulacoid,