Rhodoliths (Corallinales, Rhodophyta) As a Biosedimentary System in Arctic Environments (Svalbard Archipelago, Norway)
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Rhodoliths (Corallinales, Rhodophyta) as a Biosedimentary System in Arctic Environments (Svalbard Archipelago, Norway) Rhodolithe (Corallinales, Rhodophyta) als biosedimentäres System in der Arktis (Svalbard Archipel, Norwegen) Der Naturwissenschaftlichen Fakultät der Friedrich-Alexander-Universität Erlangen-Nürnberg zur Erlangung des Doktorgrades Dr. rer. nat. vorgelegt von Sebastian Teichert aus Nürnberg Als Dissertation genehmigt von der Naturwissenschaftlichen Fakultät der Friedrich-Alexander-Universität Erlangen-Nürnberg Tag der mündlichen Prüfung: 18. März 2013 Vorsitzender der Promotionskommission: Prof. Dr. Johannes Barth Erstberichterstatter: Prof. Dr. André Freiwald Zweitberichterstatter: PD Dr. Axel Munnecke Gaudeamus igitur iuvenes dum sumus Kindleben (1781) Content Acknowledgements .................................................................................................. I Abbreviations ............................................................................................................ III Summary ....................................................................................................................... IV Zusammenfassung ..................................................................................................... V 1. Aim of the study ................................................................................................. 1 2. Study area .......................................................................................................... 3 2.1 Geographical setting ..................................................................................................... 3 2.2 Climatic and hydrographic conditions ....................................................................... 4 2.3 Topography ..................................................................................................................... 5 2.4 Environmental characteristics of the study sites ........................................................ 6 3. Introduction to red algae (Rhodophyta) .......................................................... 9 3.1 Taxonomy of the Rhodophyta ................................................................................... 9 3.2 Species numbers and classification of the Rhodophyta ........................................... 10 3.3 Biological characteristics of the Rhodophyta ............................................................ 11 3.4 Phylogeny and evolutionary relationships of the Rhodophyta .............................. 12 3.5 Geological record and evolution of the Corallinales ............................................... 13 3.6 Environmental adaptations of the Corallinales ........................................................ 16 4. Introduction to rhodoliths ................................................................................ 20 4.1 Definition, shape, and morphology of rhodoliths ................................................... 20 4.2 Biomineralisation and growth mechanisms of rhodoliths ..................................... 21 4.3 Geological, palaeontological, and climatological significance of rhodoliths ........ 24 4.4 Geographical distribution of recent rhodolith beds ................................................ 25 5. Introduction to cool-water carbonates ............................................................. 27 5.1 A short history of cool-water carbonate research ...................................................... 27 5.2 Definition of cool-water carbonates ............................................................................ 27 5.3 Controls on cool-water carbonates .............................................................................. 27 5.4 Cool-water carbonates in the Arctic ............................................................................ 29 6. Material and methods ....................................................................................... 30 6.1 Overview ........................................................................................................................ 30 6.2 Exploration of the seafloor ........................................................................................... 30 6.3 Sampling of the water column ..................................................................................... 30 6.4 Measurements of the light regime ............................................................................... 32 6.5 Sampling of the benthic community ........................................................................... 32 6.6 Sampling of the rhodoliths ........................................................................................... 32 6.7 Taxonomy of the coralline algae .................................................................................. 33 6.8 Description and classification of the rhodoliths ........................................................ 34 6.9 Computer tomography scans of the rhodoliths ......................................................... 34 6.10 Verification of the annual rhodolith banding pattern .............................................. 35 6.11 Calculation of the annual carbonate production by the rhodolith beds ............... 35 7. Results ................................................................................................................ 37 7.1 Seafloor topography and distribution of rhodolith beds ......................................... 37 7.2 Temperature and salinity patterns ............................................................................... 48 7.3 The light regime ............................................................................................................. 51 7.4 Chemistry of the seawater ............................................................................................ 53 7.5 Composition of the benthic community .................................................................... 54 7.6 Taxonomy of the coralline algae ................................................................................. 62 7.7 Description and classification of the rhodoliths ........................................................ 67 7.8 Verification of the annual rhodolith banding pattern .............................................. 70 7.9 The annual carbonate production by the rhodoliths ............................................... 72 8. Discussion ......................................................................................................... 77 8.1 Environmental controls on Arctic rhodoliths ............................................................. 77 8.2 Interactions between rhodoliths and other benthic organisms .............................. 88 8.3 Rhodolith carbonate production ................................................................................ 91 8.4 Comparison with other rhodolith communities ....................................................... 97 8.5 Future implications ....................................................................................................... 99 9. Conclusions ....................................................................................................... 101 References ...................................................................................................................... 102 Online data sources ....................................................................................................... 117 Appendix A Raw data CTD-measurements Appendix B Raw data PAR-measurements Appendix C Raw data rhodolith size- and morphology-measurements Appendix D Raw data rhodolith surface- and protuberance-measuremens Appendix E Raw data protuberance weight- and increment-measurements Appendix F Article citation concerning the Nordkappbukta rhodolith communi- ties published in the ISI journal Phycologia Acknowledgements At this point, I would like to take the chance to express my gratitude to the persons that kindly accompanied me during my work. I greatly acknowledge my doctoral thesis supervisor, Prof. Dr. André Freiwald (Wilhelms- haven), for encouraging me to start this interesting project, his permanently helpful guidance, and for the possibility to work under such excellent conditions as I came upon them. In this coherence, I would like to acknowledge the Deutsche Forschungsgemeinschaft, for funding this work within the project FR 1134/18. I am also very thankful to PD Dr. Axel Munnecke (Erlangen), who kindly agreed to act as my second supervisor and shared his large expertises in fruitful discussions. The RV Maria S. Merian expedition this study is based on took place in the year 2006. At that time, I just had finished my basic studies and did not participate on the cruise. A fortiori, I am indebted to the members of the crew of RV Maria S. Merian and the attendant scientists for the sampling and data acquisition, which are the groundwork of this study. Some of the findings from my work have already been published in an article in the jour- nal Phycologia, dealing with the northernmost rhodolith communities known so far. Being my first peer-reviewed publication, I was very happy with the helpful comments, the criti- cal suggestions, and the substantial legwork from my co-authors, Dr. William Woelkerling (Bundoora), Dr. Andres Rüggeberg (Gent), Dr. Max Wisshak (Wilhelmshaven), Prof. Dr. Dieter Piepenburg (Kiel), Dr. Michael Meyerhöfer (Kiel), Dr. Armin Form (Kiel), Dipl.-Geol. Jan Büdenbender (Kiel), and Prof. Dr. André Freiwald. In this context, I