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Diversity and Morphology of Calcareous Dinophytes (Thoracosphaeraceae, Peridiniales)

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Diversity and Morphology of Calcareous Dinophytes (Thoracosphaeraceae, Peridiniales) Dissertation zur Erlangung des Doktorgrades der Naturwissenschaften an der Fakultät für Biologie der Ludwig-Maximilians-Universität München Diversity and morphology of calcareous dinophytes (Thoracosphaeraceae, Peridiniales) Carmen Zinßmeister München, Februar 2013 Erklärung Diese Dissertation wurde im Sinne von §12 der Promotionsordnung von PD Dr. Marc Gottschling betreut. Ich erkläre hiermit, dass die Dissertation nicht einer anderen Prüfungskommission vorgelegt worden ist, und dass ich mich nicht anderweitig einer Doktorprüfung ohne Erfolg unterzogen habe. Eidesstattliche Erklärung Ich versichere hiermit an Eides statt, dass die vorgelegte Dissertation von mir selbstständig und ohne unerlaubte Hilfe angefertigt wurde. Carmen Zinßmeister 1. Gutachter: PD Dr. Marc Gottschling 2. Gutachter: Prof. Dr. Susanne Renner Dissertation eingereicht am: 11.02.2013 Mündliche Prüfung am: 26.04.2013 i ii Note This doctoral research project has been carried out under the supervision of PD Dr. Marc Gottschling at the Ludwig-Maximilians-University Munich from February 2009 until now. The project was funded by Deutsche Forschungsgemeinschaft (DFG) in cooperation with Prof. Dr. Helmut Keupp, geological science; department paleontology at the FU Berlin and Prof. Dr. Helmut Willems, historical geology and palaontology at the University of Bremen. This thesis is based on and includes the following articles / manuscripts: Chapter 1 Who am I – and if so, how many? Species diversity of calcareous dinophytes (Thoracosphaeraceae, Dinophyceae) in the Mediterranean Sea. Soehner S., C. Zinssmeister, M. Kirsch, M. Gottschling. Org Divers Evol 12(4): 339-348 Own contribution: Field work (40%); strain isolation and cultivation (50%); morphological analysis (incl. images: 100%); manuscript preparation (20%). Chapter 2 Same but different: Two novel bicarinate species of extant calcareous dinophytes (Thoracosphaeraceae, Dinophyceae) from the Mediterranean Sea. Zinssmeister, C., S. Soehner, M. Kirsch, E. Facher, K. J. S. Meier, H. Keupp, M. Gottschling J Phycol. 48(5): 1107-1118 Own contribution: Field work (40%); cultivation (50%); species descriptions and images (90%); manuscript preparation (30%). iii Chapter 3 Catch me if you can: the taxonomic identity of Scrippsiella trochoidea (F.Stein) A.R.Loebl. (Thoracosphaeraceae, Dinophyceae). Zinssmeister, C., S. Soehner, E. Facher, M. Kirsch, K. J. S. Meier, M. Gottschling (2011). Syst Biodivers 9(2): 145-157. Own contribution: Field work (40%); strain isolation and cultivation (60%); morphological analysis (incl. images: 90%); manuscript preparation (50%). Chapter 4 Delimitation of the Thoracosphaeraceae (Dinophyceae), including the calcareous dinoflagellates, based on large amounts of ribosomal RNA sequence data. Gottschling, M., S. Soehner, C. Zinssmeister, U. John, J. Plötner, M. Schweikert, K. Aligizaki, M. Elbrächter (2012). Protist 163(1): 15-24. Own contribution: cultivation (25%); morphological analysis and images (80%); manuscript preparation – morphological part (80%). Chapter 5 Ultrastructure of calcareous dinophytes (Thoracosphaeraceae, Peridiniales) with a focus on vacuolar crystal-like particles. Zinssmeister, C., H. Keupp, G.Tischendorf, F. Kaulbars, M. Gottschling (2013). PlosONE 8(1): e54038. Own contribution: cultivation (50%); morphological, anatomical analysis and images (80%); manuscript preparation (40%). iv Summary Dinophytes are unicellular eukaryotic algae that, together with their closest relatives, ciliates and apicomplexans, belong to the superphylum of alveolates. Some of them, namely calcareous dinophytes (Thoracosphaeraceae, Peridiniales), develop an immotile calcareous cell during their life history. They accumulate in the oceans’ sediments analogously to terrestrial seed banks. Although the diversity of calcareous dinophytes was investigated in several studies, only a few of them provide data from coastal waters and sediments. The main goal of this thesis was to record the diversity of calcareous dinophytes from marine environments using morphological, anatomical, taxonomical and evolutionary approaches. An essential part of the project was to establish living dinophyte cultures, assuring constant access to fresh material for morphological and molecular analysis (chapters 1-5). The morphological diversity of extant species as well as those described from the fossil record was documented (chapter 1), and the conflict between molecular and the morphological data was presented (chapter 1-5). Furthermore, detailed morphological descriptions of two new species, Scrippsiella bicarinata und S. kirschiae (chapter 2) and morphological analysis of Bysmatrum sp., a species of doubtful phylogenetic position (chapter 4), were provided. Scrippsiella trochoidea (basionym: Glenodinium trochoideum), a species with a previously ambiguous description, has been redescribed and epitypified by myself based on material collected from the type locality to assure a reliable determination of this species (Chapter 3). Comparative ultrastructure investigations using light and electron microscopic techniques at various stages of the life cycle showed that the anatomical structure during the biomineralization processes differs within subgroups of calcareous dinophytes (chapter 5) and could be used as a useful phylogenetic trait. v Zusammenfassung Dinophyten sind einzellige Eukaryoten und bilden zusammen mit ihren nächsten Verwandten, den Ciliaten und Apicomplexa, das Taxon der Alveolaten. Innerhalb der Dinophyten besitzen Vertreter der kalkigen Dinophyten (Thoracosphaeraceae) die Fähigkeit, Kalkstrukturen während ihres Lebenszyklus‘ auszubilden, die sich in marinen Sedimenten anreichern können. Obwohl sich mehrere Studien mit der Erfassung der Diversität bei kalkigen Dinophyten beschäftigen, waren küstennahe Bereiche bisher wenig erforscht. Ziel dieser Arbeit ist die geographisch engmaschige Aufsammlung von Proben mithilfe eines Schwerelots, das dies in kürzester Zeit ermöglicht. Im Rahmen der vorliegenden Dissertation wurde die Diversität kalkiger Dinophyten morphologisch und anatomisch mithilfe von licht- und elektronenmikroskopischen Methoden untersucht. Die Grundlage hierfür bildete die Etablierung einer Lebendsammlung von Dinophyten, die den Zugriff auf frisches Material für morphologische und molekulare Analysemethoden gewährleistete (Kapitel 1-5). Hierdurch gelang es eine Vielzahl rezenter und auf fossilen Beschreibungen basierender Arten (Kapitel 1) kalkiger Dinophyten zu kultivieren, deren morphologische Vielfalt darzustellen und diese molekular-phylogenetischen Daten gegenüber zu stellen (Kapitel 1-4). Dies umfasste außerdem die detaillierte morphologische Beschreibung zweier neuer Arten, Scrippsiella bicarinata und S. kirschiae, (Kapitel 2), morphologische Analysen zur Klärung der systematisch problematischen Stellung von Bysmatrum sp. (Kapitel 4) und die Kultivierung und Untersuchung von kryptischen (morphologisch nicht abgrenzbaren) Scrippsiella cf. trochoidea Stämmen (Kapitel 1, 3). Aufgrund von unzureichend fixiertem, dokumentiertem oder nicht vorhandenem Typusmaterial stellte eine präzise Artidentifizierung mitunter eine Herausforderung dar. Durch Epitypisierung gelang es, den wissenschaftlichen Namen Scrippsiella trochoidea (Basionym: Glenodinium trochoideum) taxonomisch zu klären und in Zukunft eine verlässliche Bestimmung dieser Art zu ermöglichen. (Kapitel 3). Des Weiteren zeigten vergleichende Ergebnisse von ultrastrukturellen Untersuchungen während der Biomineralisation von kalkigen Dinophyten, dass sich die Kalzifizierungs-Prozesse innerhalb der Subgruppen unterscheiden (Kapitel 5) und teilweise als phylogenetisches Merkmal genutzt werden können. vi Content Erklärung ..................................................................................................................................... i Ehrenwörtliche Versicherung ...................................................................................................... i Note ....................................................................................................................................... iii Summary .................................................................................................................................... v Zusammenfassung ..................................................................................................................... vi Content ..................................................................................................................................... vii Introduction ................................................................................................................................ 1 Biodiversity and taxonomic research ...................................................................................... 1 Life style and ecology of calcareous dinophytes (Thoracosphaeraceae) ................................ 3 Phylogeny and classification of calcareous dinophytes .......................................................... 4 Morphology and anatomy of calcareous dinophytes .............................................................. 7 Morphology of the motile theca .......................................................................................... 7 Morphology of the calcareous coccoid cell ...................................................................... 10 Biomineralization within calcareous dinophytes .............................................................. 12 References ............................................................................................................................
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