Rhodoliths (Corallinales, Rhodophyta) As a Biosedimentary System in Arctic Environments (Svalbard Archipelago, Norway)

Home , Lithophylloideae, Lithothamnion, Lithothamnion glaciale, Rhodolith, Sporolithaceae

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 communities 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 journal 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 critical 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 furtherly express my great gratitude to Dr. William Woelkerling, who accompanied me during my PhD period with his great knowledge and experience on red algae and scientific writing. For being on the spot all the time, I am deeply indebted to him.

The GeoZentrum Nordbayern in Erlangen is the place where I conducted my research, and the staff of the Institute for Palaeontology has always afforded a very contented time to me. I greatly acknowledge Petra Wenninger for finding safe paths in the jungles of bureaucracy, Fritz Ruhmann for giving a helpful hand whenever needed, and Birgit Leipner-Mata for help and advice in the laboratory.

I I also enjoyed the time I spent with my dear colleague Dipl.-Geol. Matthias López Correa (Erlangen), having discussions about science and life in general, or just going rock climbing. Thank you for the good times inside and outside the institute.

Special thanks go to Dipl.-Geol. Christian Schulbert (Erlangen), who ministered to me from my first day at the institute. He accompanied me during my diploma thesis and my PhD- period, and I am happy to have him as a colleague and as a friend.

Already as a child, I was interested in nature, minerals, and fossils. And from that time on, there have been people that encouraged me and kept those interests alive. These people were Alois de Pauli, Siegfried Nitsch, and my father, and I am thankful that they did so.

From a private point of view, I would like to thank my dear friends Friederike Adomat, Andreas Wässerle, and especially Ines Pyko, for their support, their encouragement, and all the great moments I was able to share with them.

Finally, my dearest thanks go to my parents, and to enumerate all the things they did for me, would go beyond the scope of this chapter, for sure.

II Abbreviations

a: (Lat. annum) Jahr mA: milliampere Arg: aragonite MB: multibeam ArW: Arctic Water n: n molar solution AW: Atlantic Water n.d.: no data c.: (Lat. circa) about ΩArg: aragonite saturation ca.: (Lat. circa) ungefähr ΩCal: calcite saturation Cal: calcite p: pressure cf.: (Lat. conferre) confer! p.: page CAC: coralline algal crusts pp.: ‘plural of page’ CTD: conductivity, temperature, and depth PAR: photosynthetic active radiation dbar: decibar pCO2: CO2-partial pressure DIC: dissolved inorganic carbon PSS: practical salinity scale DRG: dredge SEM: scanning electron microscope EDX: energy dispersive X-ray stat #: station number ESC: East Spitsbergen Current SW: seawater IPCC: Intergovernmental Panel on Climate Change T: temperature kHz: kilohertz TA: total alkalinity Lat.: latitude WSC: West Spitsbergen Current Long.: longitude yr: year LS: light measurement

III Summary

Polar coralline algae (Corallinales, Rhodophyta) that form rhodoliths have received little at- tention concerning their potential as carbonate factories. This study is the first attempt to quantify the carbonate production by rhodolith beds from four settings along the coast of Svalbard, situated at different latitudes, as well as to assess their environmental controls and their role as autogenic ecosystem engineers sensu Jones et al. (1994), meaning that they create habitats by modifying themselves. The settings in order from south to north are Floskjeret (78°18’N), Krossfjorden (79°08’N), Mosselbukta (79°53’N), and Nordkappbukta (80°31’N), featuring the northernmost rhodolith communities currently known. That far beyond the polar circle, the perennial algae must be adapted to extreme seasonal variations in light regime (c. 4 months of winter darkness), temperature, salinity, and sea ice coverage. During the MSM 02/03 expedition of RV Maria S. Merian in 2006, the rhodolith communities and their environment were investigated using multibeam swath bathymetry, CTD measurements, record- ings of the photosynthetic active radiation, determinations of the water chemistry, seabed imaging and targeted sampling by means of the manned submersible JAGO, as well as benthic organism collections with a dredge. The carbonate production of the rhodoliths was calculated from fuchsine stained annual rhodolith growth increments and the density of the protuberances in the rhodolith beds. The results were verified using statistical methods and Mg/Ca ratio EDX-measurements. The rhodolith communities are mainly composed of Lithotham- nion glaciale Kjellman, 1883, with a lesser amount of Phymatolithon tenue (Rosenvinge) Düwel & Wegeberg, 1996. Most rhodoliths are monospecific, consisting either of L. glaciale or P. tenue. The rhodolith shape and size varies considerably among the particular sites in relation to the colonised lithoclasts. Rhodoliths occur between 47 and 27 m water depth, while coralline algal crusts on lithoclastic cobbles appear as deep as 77 m water depth. They display a patchy distribution and different stages of development, ranging from initial crusts to well developed rhodoliths. The carbonate production rates range from 100.9 g m-2 yr-1 at Nordkap- pbukta to 200.3 g m-2 yr-1 at Floskjeret, and significantly depend on the degree of latitude. The study shows that polar rhodoliths are important carbonate producers and ecosystem engineers, and are much more widespread in Arctic waters than previously thought. At each site, the ambient waters are saturated with respect to both aragonite and calcite, and the rhodolith beds are generally located at dysphotic depths (<1% surface illumination). The rhodolith- associated macrobenthic fauna features prominent grazers like chitons and sea urchins, which keep the coralline algae free from epiphytes. Overall, L. glaciale and P. tenue are well adapted to the extreme environment of the Arctic. The results of this study provide a better understanding of the relationships between rhodolith communities, their environment, and their susceptibility to climate change.

IV Zusammenfassung

Polare, coralline Rotalgen (Corallinales, Rhodophyta), die Rhodolithe bilden, haben bislang wenig Aufmerksamkeit in Bezug auf ihr Potential als Ökosystem-Bildner und als Karbonat- fabriken erhalten. Diese Studie ist der erste Versuch, sowohl die Karbonatproduktion durch Rhodolithvorkommen vier verschiedener Lokalitäten von unterschiedlichen Breitengraden vor der Küste Svalbards zu quantifizieren. Sie zielt auch darauf ab, ihre Abhängigkeiten von Um- weltfaktoren und ihre Rolle als autogene Ökosystem-Bildner sensu Jones et al. (1994; d.h. sie erzeugen Habitate indem sie sich selbst modifizieren) zu beurteilen. Die Lokalitäten von Nord nach Süd sind Floskjeret (78°18‘N), Krossfjorden (79°08‘N), Mosselbukta (79°53‘N) und Nord- kappbukta (80°31`N), wo das bislang nördlichste Rhodolithvorkommen entdeckt wurde. So weit jenseits des Polarkreises müssen die mehrjährigen Algen an die extreme Saisonalität des Licht- klimas (ca. 4 Monate Polarnacht), der Temperatur, der Salinität und der Eisbedeckung angepasst sein. Im Verlauf der MSM 02/03 Expedition der RV Maria S. Merian im Jahr 2006 wurden die Rhodolithgemeinschaften und ihre Umwelt mit einem Multibeam-Fächerecholot, CTD-Mes- sungen, Messungen der photosynthetisch wirksamen Strahlung, Bestimmungen des Wasserche- mismus, der Dokumentation und gezielten Beprobung des Meeresbodens mit dem bemannten Tauchboot JAGO, sowie durch die Sammlung benthischer Organismen mit einer Dredge er- forscht. Die Karbonatproduktion der Rhodolithe wurde durch die Auszählung Fuchsin-gefärb- ter Rhodolith-Wachstumsinkremente in Kombination mit der Dichte der Protuberanzen in den Rhodolithakkumulationen berechnet. Die Ergebnisse wurden durch statistische Methoden und EDX-Messungen der Mg/Ca-Ratio verifiziert. Die Rhodolithgemeinschaften werden hauptsäch- lich von Lithothamnion glaciale Kjellman, 1883 und zu einem geringeren Prozentsatz von Phy- matolithon tenue (Rosenvinge) Düwel & Wegeberg, 1996 gebildet. Die meisten Rhodolithe sind monospezifisch und bestehen entweder aus L. glaciale oder P. tenue. Die Form und Größe der Rhodolithe variiert stark zwischen den einzelnen Lokationen und ist abhängig von den be- siedelten Lithoklasten. Rhodolithe kommen zwischen 27 und 47 m Wassertiefe vor, während Krusten coralliner Rotalgen auf Lithoklasten bis in 77 m Wassertiefe zu finden sind. Sie zeigen eine fleckenhafte Verteilung und unterschiedliche Entwicklungsstufen, angefangen von initialen Krustenstadien bis hin zu gut entwickelten Rhodolithen. Die Karbonatproduktion reicht von 100,9 g m-2 a-1 in Nordkappbukta bis zu 200,3 g m-2 a-1 in Floskjeret und hängt stark vom jeweili- gen Breitengrad ab. Die Studie zeigt, dass polare Rhodolithe wichtige Karbonatproduzenten und Ökosystem-Ingenieure sind, deren Verbreitung in arktischen Gewässern bislang unterschätzt wurde. Das Wasser jeder Lokalität ist gesättigt mit Kalzit und Aragonit und die Rhodolithak- kumulationen befinden sich hauptsächlich in der dysphotischen Zone (<1% der Oberflächen- Lichtmenge). Die makrobenthische Begleitfauna enthält bekannte Weidegänger wie Käferschne- cken und Seeigel, welche die Rhodolithen frei von Epiphyten halten. Insgesamt sind L. glaciale und P. tenue sehr gut an die extremen Umweltbedingungen der Arktis angepasst. Die Ergebnisse dieser Studie tragen zu einem besseren Verständis der Beziehungen zwischen Rhodolithgemein- schaften, ihrer Umwelt und ihrer Anfälligkeit für klimatische Veränderungen bei.

V Aim of the study

1. Aim of the study

The description and interpretation of the nearly infinite variety of carbonate rocks and sediments have occupied geologists for more than two centuries (Pomar & Hallock 2008). The depositional features vary in space and time, and Schlager (2000; 2003) defined, according to the sedimentation mode, three benthic carbonate factories, being the tropical shallow-water factory, the cool-water factory, and the mud-mound factory. The cool-water carbonates were largely ignored until the 1980s, the so-called golden age of carbonate research (James 1997), and were also regarded to be dominated by heterotrophic skeletal production (Schlager 2000). The least known cool-water carbonate factories until now are those of the polar region (Ro- gala et al. 2007). In Nordic polar environments, the open shelf Spitsbergen Bank and shelf banks southwest off Spitsbergen were known as carbonate factories driven predominantly by molluscs and barnacles (Andruleit et al. 1996; Henrich et al. 1997), and findings by Frei- wald (1993) and Freiwald & Henrich (1994) showed that also photoautotrophic coralline algae that form rhodoliths play a major role as a cool-water carbonate factories than estimated before. In 2006, the northernmost rhodolith communities so far known were discovered at 80°31’N at Nordkappbukta (Nordaustlandet, Svalbard) during the MSM 02/03 expedition of RV Maria S. Merian (see Lherminier et al. 2006 for the cruise report). This and three other sites around the Svalbard archipelago, Floskjeret in Isfjorden, Krossfjorden in the Kongsfjord-Krossfjorden system, and Mosselbukta at the northernmost tip of Ny-Friesland were intensively sampled and documented. Already during the cruise, the preliminary results showed that coralline algae are very widespread in that polar and extremely seasonal environment. The present study analyses the biotic assemblage and functioning of this poorly known biosedimentary system and calculates the annual carbonate productions of the prevailing rhodolith beds in relation to environmental parameters. The geographic position of Svalbard exposes the archipelago to a very strong seasonality. Hence, the coralline algae thriving at such localities have to be adapted to strong seasonal changes in temperature, salinity, nutrient levels, and the light regime. Especially the light regime including the polar night which lasts for up to 126 days at Svalbard (data from USNO Sun Rise Tables) highly stresses these photosynthetic organisms. Even if they can only record snap shot conditions, various measurements of these parameters using gear such as CTD and sensors for the photosynthetic active radiation (PAR) should help to specify the prevailing conditions and enlight the adaptations and demands of the rhodolith forming coralline algae. As mentioned above, Svalbard is strongly affected by the polar climate, and the seafloor at the investigated sites is mainly made up by glaciogenic flats and moraines (Sexton et al. 1992; Harland 1997). These flats consist of lithoclasts of pebble and cobble size while boulders larger than one cubic metre are very rare. For this reason, the lithoclasts are the only stable substratum for the growth of the coralline algae and offer little shelter and niches for the

1 Aim of the study prevailing benthic organisms. Dredge samples have been used to show the composition and the functional organism groups at the four sites, and the JAGO video footage has been used to determine the biotic interactions between the epibenthic organisms and the coralline algal crusts and rhodoliths. This includes the role of the rhodoliths as autogenic bioengineering organisms sensu Jones et al. (1994), meaning that they create habitats by their own skeletal growth and the successive modification of this skeleton. The video footage has also been used to distinguish the required features of a substratum to be colonised by coralline algae. Since the coralline algae produce a massive calcitic skeletal framework, their role as a potential carbonate factory has yet not been considered. More than 300 individual rhodoliths were collected during the cruise in order to estimate the carbonate production per square metre per year at a particular site. The obtained data have been used to detect the possibly influencing parameters on carbonate production and to discuss the possible consequences of the ongoing climate change. Overall, the present study provides a better understanding of the rhodolith forming coralline algae from Svalbard and their susceptibility to environmental parameters. It underpins their role as a hitherto underestimated biosedimentary system of the Arctic.

2 Study area

2. Study area

2.1 Geographical setting

Svalbard is an archipelago, which 10°E 15°E 20°E 25°E is situated on the north-western

margin of the Barents Shelf, 81°N 1 = Floskjeret 650 km north of the mainland of 2 = Kross orden 4 N Norway, and includes all islands 3 = Mosselbukta in the region between 74°-81°N 4 = Nordkappbukta and 10°-35°E. The main islands 3 are Spitsbergen, Nordaustlandet, Nordaustlandet Barentsøya, Edgeøya, and Prins Karls Forland. Spitsbergen is the Hinlopenstretet central island, Nordaustlandet 2 Forlandsundet in the northeast is divided from 79°N 80°N Spitsbergen Spitsbergen by Hinlopenstre- 1 Barentsøya tet, Barentsøya and Edgeøya in Longyearbyen the southeast are divided from Forland Karls Prins Is orden 78°N Spitsbergen by Storfjorden, and Edgeøya Stor orden Prins Karls Forland in the west is divided from Spitsbergen by For- landsundet. The study sites (Fig. 77°N 2.1) are, in order from south to north, Floskjeret in Isfjorden (78°18’N, 14°32’E), Krossfjorden Fig. 2.1. Map of the Svalbard archipelago (modified after Har- land 1997) with the main islands Spitsbergen, Nordaustlandet, in the Kongsfjorden-Krossfjor- Barentsøya, Edgeøya, and Prins Karls Forland; the yellow stars den system (79°08’N, 11°40’E), indicate the four study sites. Mosselbukta at the northernmost tip of Ny-Friesland (79°53’N, 15°55’E), and Nordkappbukta at the northernmost tip of Nordaustlandet (80°31’N, 19°52’E). The name ‘Svalbard’, which means ‘cool coast’, was formally introduced by Orvin (1942) and since 1920, the Spitsbergen Treaty recognises the absolute sovereignty of Norway over the archipelago. The first permanent communities were established at the beginning of the 20th century with the startup of coal mining (Hoel 1966). Today, the administrative centre of Sval- bard is Longyearbyen and other settlements are the Russian mining community of Barents- burg and the research community of Ny-Ålesund. Svalbard’s present-day population amounts to c. 2400, most of which (c. 2100) are living in Longyearbyen (data from Statistics Norway).

3 Study area

2.2 Climatic and hydrographic conditions

The polar regions are characterized by pronounced seasonal variations of the light regime, low temperatures, and long periods of snow and ice cover (Wiencke et al. 2007). Since the Svalbard archipelago is situated between 74°-81°N, many distinctive attributes of this Arctic environment derive from the angle of incidence of the solar radiation, e.g. the duration of each the polar day, i.e. the sun stays above the horizon for >24 h, and the polar night, i.e. the sun stays below the horizon for >24 h (Kosak 1967). These phenomena occur only inside the polar circles (66°33’44’’), and Table 2.1 clearly shows that the duration of the polar day and the polar night increases with the geographical latitude, resulting in a Table 2.1. Duration of the polar day and night strong seasonality of the light regime. depending on the geographical latitude (data The climate of the relatively small Svalbard from USNO Sun Rise Tables). archipelago is influenced by two sources of Geogr. Number of days of continuous surface ocean water (Fig. 2.2). One is the Latitude Daylight Darkness West Spitsbergen Current (WSC), which is a 80.0°N 137 123 relatively warm and saline water mass, being 75.0°N 131 109 the northernmost extension of the Norwe- 70.0°N 72 53 gian Atlantic Current respectively the north- 66.5°N 17 0 ernmost remnant of the Gulf Stream (Orvik & Niiler 2002). The core temperatures of the WSC range from 6-8°C and the salinities from 35.1-35.3 (Aagaard et al. 1987). The other is the the polar East Spitsbergen Current (ESC), which brings cold water and drift ice southwestwards east of Spitsbergen and the eastern islands (Harland 1997). The core temperatures of the ESC range from 1-3°C and the salinities from 34.5-34.9 (OSPAR Commission 2000). These currents meet off Sørkapp and the cold ESC water is deflected and continues northwards between ESC the warmer WSC water and the coast, often carrying drift ice with it (Hisdal 1985). WSC In the northern part of Svalbard (Ny-Fries- land and Gustav V Land), sea ice usually forms with the beginning of October and Fig. 2.2. Progression of the warm, atlantic WSC starts to break up between July and August (red) and the cold, polar ESC (blue) along the (Falk-Petersen et al. 2000; Spreen et al. coasts of Svalbard; the yellow star indicates the 2008; data from AMSR-E Sea Ice Maps). In position of SØrkapp (modified after Svendsen et the western part of Svalbard (Haakon VII al. 2002). Land and Oscar II Land), sea ice usually

4 Study area forms with the beginning of January and starts to break up between April and July (Węslawski et al. 1995; Svendsen et al. 2002; Nilsen et al. 2008; Spreen et al. 2008; data from AMSR-E Sea Ice Maps). The tides range between c. 2 m for spring tide and c. 1 m for neap tide if there is no restriction by land (Hisdal 1985). The annual precipitation is low, averaging c. 200-300 mm on the western coast with a maximum of c. 1000 mm (Hisdal 1985), most of which falls as fine snow or rain in summer and autumn. The air temperature at sea level averages c. 4-5°C in summer and c. -10°C in winter. The temperatures are lower towards the north and east and the extreme range is from -50°C to 20°C (Hisdal 1985; Harland 1997).

2.3 Topography

The Central Basin of Svalbard was formed around the time of the Cretaceous-Tertiary boundary, during the strike-slip-dominated movements of Greenland and the Barents Sea Block (Spielhagen & Tripati 2009). The Barents Shelf belongs to the northwestern margin of Eu- ropes continental lithosphere and a extends northwards from Nor- way and northwestern Russia. Ex- cept for the northwestern margin, where the Svalbard archipelago and Franz Josef Land emerge, the shelf is covered by the Barents Sea, and the northern margin of the shelf is marked by the continental slope down to the Polar Ocean Basin (Harland 1997). The northern shelf bathymetry b reflects a Neogene and Quater- nary history as well, showing a subdued drainage pattern. The southwestern shelf is a pas- sive continental margin, rifted and sheared during the Tertiary (Müller & Spielhagen 1990). During the Weichselian glaciation, the southwestern shelf was Fig. 2.3. Typical landforms sculptured by glacial erosion at the iceloaded several times to alter- (a) northeastern and (b) western coast of Spitsbergen (images nating glacier advances and melt- provided by the participants of the MSM 02/03 cruise). downs (Andruleit et al. 1996).

5 Study area

The topography of mainland Svalbard reflects the geological structures which determine many contrasting landforms. Repeated sea-level changes have eroded and then exposed large tracts of nearly flat land and the land sculpture is a continuation of glacial erosion, resulting in steep cliffs, valleys, and glaciers (Harland 1997; see also Fig. 2.3). The mountain peaks fall within a general summit envelope that represents an uplifted and warped peneplain almost regardless of the altitude of the strata. The mountain contours result from a cold-desert environment, with steep slopes and cliffs where resistant rocks prevail, so the opportunity for the establishment of vegetation is strongly limited, and a more subdued landscape made up by soft rock (Harland 1997). This variety of landforms is typical for the Arctic (Thorén 1969). The coastal topography is dominated by large, glacially eroded fjords. This holds particularly for the western and the northern coasts of the island Spitsbergen, where Isfjorden and Wijde- fjorden have lengths of >100 km (Hisdal 1985; Harland 1997). Where it is allowed by the relief, the coastal areas do also exhibit strandflat topography with low-lying plains (Hisdal 1985) and the coastal sediments are strongly affected by glacial activity (Harland 1997). Since the forming mechanisms took place at a large scale, the characteristics described above apply for the Svalbard archipelago in general. In the following section, the small scale differences of the study sites examined will be presented.

2.4 Environmental characteristics of the study sites

The MSM 02/03 expedition of RV Maria S. Merian was carried out in the year 2006 and led around Spitsbergen and along the northern coast of Nordaustlandet (see Lherminier et al. 2006 for the cruise report). Because of the various research interests of the participating institutions, a great many places were visited. The sites in the focus of this study are characterised by the presence of extensive rhodolith beds and are located around Spitsbergen and Nordaustlandet (Fig. 2.4a).

Floskjeret (78°18’N, 14°32’E; Fig. 2.4b) is located at the mouth of Borebukta in Isfjorden. Isfjorden is the largest fjord system on Spitsbergen (c. 100 km long and up to 425 m deep), consisting of the trunk fjord Isfjorden and 13 tributary fjords and bays (Forwick & Vorren 2009). The geology features Early Cretaceous sediments mainly consisting of sandstones from the Hel- vetiafjellet and Carolinefjellet formations (Harland 1997). Water masses of internal and external origin affect the hydrography of this estuarine system. Inside the fjord, surface waters consisting of glacial melt water and river runoff during summer, local waters characterised by increased salinity due to sea ice formation, and winter-cooled waters prevail (Forwick & Vorren 2009). Atlantic Water (AW) from the WSC and Arctic Water (ArW), which is an extension of the polar ESC, enter the fjord system through the mouth of Isfjorden (Nilsen et al. 2008). Sea ice usually forms in December/January and starts to break up between April and July (Węs- lawski et al. 1995; Svendsen et al. 2002; Nilsen et al. 2008; data from AMSR-E Sea Ice Maps). The mean water temperatures are 2.1°C at the surface and 1.5°C at 50 m water depth. The mean salinities are 34.9 at the surface and 35.0 at 50 m water depth (data from LEVITUS 94).

6 Study area

Krossfjorden (79°08’N, 11°40’E; Fig. 2.4c) is oriented north to south and consists of a sub- marine channel (c. 30 km long and up to 370 m deep), which converges with Kongsfjorden in the Kongsfjordrenna, a deep glacial basin (Sexton et al. 1992; Svendsen et al. 2002). The northern Kongsfjordrenna is dominated by Mitragrunnen Bank, which is as shallow as 30 m and is separated from the main shelf area by a north-south oriented trough (Ottesen et al. 2007). The Kong Haakons Halvøya divides the inner fjord into two parts, Möllerfjorden and Lilliehöökfjorden. The geology features Late Neoproterozoic basement consisting of pelites from the Signeham- na formation and granites from the Generalfjella formation (Harland 1997). Seasonal variations in freshwater input are responsible for a stable stratification in summer, when the upper layer circulation is confined to a shallow surface layer, and a very weak stratification in winter (Svendsen et al. 2002). Especially Lilliehöökfjorden is strongly influenced by the tidewater glacier Lilliehöökbreen (Svendsen et al. 2002). The exchanged coastal water

a e b c

d e

Fig. 2.4. The study areas (red hachures) around Svalbard were extensive rhodolith beds were investigated; (a) map of Svalbard with yellow stars showing the positions of all sites; (b) Floskjeret in Isfjor- den; (c) Krossfjorden at the western coast of Spitsbergen; (d) Mosselbukta at the mouth of Wijdefjorden; (e) Nordkappbukta at the northern coast of Nordaustlandet; scale bars = 5 km; modified after topogra- phic map of Svalbard, 1:1000000.

7 Study area contains AW with increased temperature and salinity. Another water mass, the low temperature (<1°C) Local Water, spreads mostly over the AW. The surface Water is also characterised by a decreased salinity, 30 in the middle of the fjord and <28 in the inner basin near the glaciers. It is already regarded as brackish water and occupies several meters of the upper water column with a decreasing thickness towards the fjord mouth (Svendsen et al. 2002). Sea ice usually forms in January/February and starts to break up between April and July (Svendsen et al. 2002; Spreen et al. 2008; data from AMSR-E Sea Ice Maps). The mean water temperatures are 0.6°C at the surface and 0.7°C at 50 m water depth. The mean salinities are 33.6 at the surface and 34.7 at 50 m water depth (data from LEVITUS 94).

Mosselbukta (79°53’N, 15°55’E; Fig. 2.4d) is a classic locality studied by Kjellman (1883; 1885), who found rhodolith beds covering the whole seabed in some areas. The site is located next to a bay at the northeastern end of Ny-Friesland and terminates the peninsula Mossel- halvøya, being the seaward end of the great Ny-Friesland peninsula that borders Spitsbergen to the northeast and separates Wijdefjorden from Hinlopenstretet. The geology features Late Palaeoproterozoic basement of the Stubendorffbreen supergroup, mainly consisting of psammites, quartzites, and amphibolites (Harland 1997). In the area of Mosselbukta and Wijdefjorden, a mixing of AW and ArW occurs. The impact of AW is only pronounced during a period of strong activity of the WSC; during the rest of the time, the influence of ArW prevails (Sapota et al. 2009). Sea ice usually forms in December/January and starts to break up between May and July (Spreen et al. 2008; data from AMSR-E Sea Ice Maps). The mean water temperatures are 0.3°C at the surface and 0.7°C at 50 m water depth. The mean salinities are 33.3 at the surface and 34.8 at 50 m water depth (data from LEVITUS 94).

Nordkappbukta (80°31’N, 19°52’E; Fig. 2.4e) is currently the northernmost known locality where well developed coralline algal rhodolith communities occur. It is located at the northern tip of the Laponiahalvøya, Nordaustlandet, and Beverlysundet extends 4.8 km SSE, sepa- rating the Nordaustlandet mainland from a disembarkation area around Nordkapp. Nordkappbukta is the only site with exposed bedrock observed during the underwater surveys and features Early Neoproterozoic granite from the Laponiafjellet formation (Harland 1997). The prevailing ArW is influenced by the WSC, so AW mixes with ArW, but this impact is only pronounced in periods of strong WSC activity (Sapota et al. 2009). Seasonally changing freshwater input from melting glaciers additionally contributes to the local attributes of the water column. Sea ice usually forms in October/November and starts to break up between July and August (Falk-Petersen et al. 2000; Spreen et al. 2008; data from AMSR-E Sea Ice Maps). The mean temperatures are 0.2°C at the surface and 0.5°C at 50 m water depth. The mean salinities are 33.1 at the surface and 34.8 at 50 m water depth (data from LEVITUS 94).

8 Introduction to red algae (Rhodophyta)

3. Introduction to red algae (Rhodophyta)

3.1 Taxonomy of the Rhodophyta

Regarding the generic definition of the Rhodophyta Wettstein, 1901, many of the taxonomically important characters can be recognized in both fossil and recent specimens, while recent material comprises additional morphological attributes such as developmental and staining characteristics (Bosence 1991). There are more than 10000 described species of red algae. The amount of real species in this number, however, remains an unanswered question (Woelker- ling 1990). The species treated in the present study belong to the family Corallinaceae Lamouroux, 1812 (= calcareous red algae; order Corallinales Silva & Johanson, 1986) and are non-geniculate, i.e. they lack non-calcified joints. Taxonomically, the non-geniculate Corallinaceae are widely considered to be among the most diffcult and troublesome groups of the Rhodophyta (Woelk- erling 1988), and it would go beyond the scope of this work to give a complete code of practice how to identify specimens to genus or even to species level. During the nineteenth century, the genera of the non-geniculate Corallinaceae were identified principally according to the morphology of their thalli (e.g. Kützing 1849). From today’s perspective, this is a very uncertain feature because the intraspecific variabilty is high and the growth forms largely depend on external influences like water movement (Bosence 1983a) or grazing pressure from herbivores (Steneck & Adey 1976; Steneck 1986). With the upcoming of enhanced examination tools like the scanning electron microscope (SEM) and optical microscopy techniques, the diagnostic features became much more versatile including, e.g., vegetative structures such as spore germi- nation patterns, the epithallial cell shape and occurrence, or the arrangement of spores within the sporangia (Woelkerling 1988). These short remarks make clear that the identification of the Corallinaceae to genus or species level is quite complicated and for further interest, taxonomic publications (see Woelkerling 1988, and references therein) are recommended. The correct taxonomic description of rhodoliths, which are composed of one or more coralline algal taxa, is even more complicated. Rhodolith growth forms cover a wide range of those described for crustose non-geniculate corallines, but since morphological descriptions have also suffered from the use of multiple terms for one and the same feature, a universal adoption of the form names as proposed by Woelkerling et al. (1993) would improve the communication (Foster 2001). Additionally, as discussed by Johansen (1981) and Woelkerling (1988), it can still be a problem to assign an accurate genus and species name to a particular rhodolith, may it be fossil or living. This probably will remain so until thorough regional reviews and monographs are evaluated using modern taxonomic concepts, type material (Riosmena-Rodríguez et al. 1999) and molecular genetic analyses (Bailey & Chapman 1998; Bailey 1999). This important taxonomic work may enhance the knowledge of when and where taxa occur and which environmental constraints on the distribution of living species exist (Foster 2001).

9 Introduction to red algae (Rhodophyta)

3.2 Species numbers and classification of the Rhodophyta

As mentioned above, there are more than 10000 species of red algae described, while the number of real species remains uncertain (Woelkerling 1990). The estimates vary by over 100%, including 2500 (Dring 1982), 3700 (Peterfi & Ionescu 1977), 3900 (Dixon 1973; Bold et al. 1987), 4000 (Kraft & Woelkerling 1990), 3500 to 4500 (van den Hoek & Jahns 1978), 5100 (Tappan 1980), 5200 (Dixon 1973), and 6000 (Pritchard & Bradt 1984). It is extremely difficult to make accurate estimates of species numbers if reliable species concepts scarcely exist within many genera (Woelkerling 1983; 1988; Garbary 1987), if the status of substantial numbers of inadequately described species remains unresolved (Womersley 1979; Woelkerling 1984), and if there are virtually no modern, worldwide monographs of genera containing more than a few species (Woelkerling 1990). The classification and the taxonomic distribution of species within the orders and families of the Rhodophyta are unresolved and there is no overall consensus how many orders or families should be recognized (Woelkerling 1990). Data provided by Kraft & Woelkerling (1990) on the estimated numbers of families, genera and species in one suite of orders/subclasses are summarized in Table 3.1, but the authors emphasize the composite nature of this summary, which may not coincide with those of other authors. According to Woelkerling (1990), one value of generating notional estimates of species numbers was to gain insight into both, the apparent diversity of red algae relative to other algal groups, and the relative proportion of red algae occurring in different geographic areas. Dring (1982) has calculated that 27.1% of all known species of marine plants are red algae.

Table 3.1. Estimated numbers of families, genera and species within orders/subclasses of the Rhodo- phyta according to Cole & Sheath (1990) and Kraft & Woelkerling (1990). Estimated numbers of included taxa Order/Subclass Families Genera Species Acrochaetiales Feldmann, 1953 1 1-7 100 Bangiophycidae1 Wettstein, 1901 n.d. 30 110 Batrachospermales Pueschel & Cole, 1982 3 8 80 Bonnemaisoniales Feldmann & Feldmann, 1942 2 8 25 Ceramiales Oltmanns, 1904 4 325 1500 Corallinales Silva & Johansen, 1986 1 37-40 uncertain Gelidiales Kylin, 1923 2 12 130 Gigartinales Schmitz, 1892 42 170 1120 Hildenbrandiales Pueschel & Cole, 1982 1 2 11 Nemaliales Schmitz, 1892 2 18 350 Palmariales Guiry & Irvine in Guiry, 1978 1 6 20 Rhodymeniales Schmitz in Engler, 1892 3 42 280 1 Includes: Bangiales Schmitz, 1892; Compsopogonales Skuja, 1939; Porphyridiales Kylin, 1937; Rhodochaetales Bessey, 1907.

10 Introduction to red algae (Rhodophyta)

These calculations also show that species of the red algae appear to be more diverse in benthic marine environments on a world scale than species of the brown algae (Phaeophyta) and the green algae (Chlorophyta) combined, and regional data by South (1987) and Womersley (1981; 1984; 1987) show similar patterns (see Table 3.2). Another problem regarding the classification of the red algae is that some characteristics that have been used to delimit orders and families are confined to events in the sexual cycle. Hence, they are difficult to observe and have not been confirmed to occur in a majority of the included species. The extent to which new data on the reproductive biology of red algae will affect their classification both at order and family levels is difficult to forecast but may be substantial (Woelkerling 1990).

Table 3.2. Estimated species numbers of marine Rhodophyta, Phaeophyta, and Chlorophyta occurring on a world scale or in particular regions according to Cole & Sheath (1990). Region Rhodophyta Phaeophyta Chlorophyta Reference Wettstein, 1901 Wettstein 1901 Pascher, 1914 Worldwide 2540 997 900 Dring (1982) North Atlantic Ocean 589 324 258 South (1987) Southern Australia 800 231 123 Womersley (1981; 1984; 1987)

3.3 Biological characteristics of the Rhodophyta

Red algae are presumed to share several combinations of characteristics, and as stated by Woelkerling (1990), these particular combinations do not occur in any other group of organisms and are as follows:

• Eukaryotic cells • Flagella absent • Food reserves stored principally as floridean starch (amylopectin) • Food reserves stored in cytoplasm, not in chloroplasts • Phycoerythrin, phycocyanin, and allophycocyanins as accessory pigments • Chloroplasts with non-aggregated (unstacked) thylakoids • Chloroplasts lacking external endoplasmatic reticulum

Consequently, the red algae have to be considered to constitute either a separate division in the Kingdom Plantae [variously named the Rhodophyta (Kylin 1956; Dixon 1973; Bold & Wynne 1985; Kraft & Woelkerling 1990), the Rhodophycota (Parker 1982; South

11 Introduction to red algae (Rhodophyta)

& Whittick 1987), or the Rhodophycophyta (Papenfuss 1955; Bold & Wynne 1978)] or a separate phylum in the Kingdom Protoctista [named the Rhodophyta (Margulis & Schwartz 1982)]. In more than 90% of all red algal species, chlorophyll a and several carotenoids occur along with phycoerythrin and phycocyanin (Ragan 1981), providing a much larger wavelength spectrum for photosynthesis than it was available if only chlorophyll a, chlorophyll b, and carotenoids were present (Fig. 3.1). Varying proportions of these pigments can result in red, pink, violet, blue, brown, black, yellowish, or greenish thalli. However, the most marine species are red or pink, whereas most freshwater species are bluish to black (Woelkerling 1990).

UV-A IR

350 400 450 500 550 600 650 700 750 800 nm chl-a chl-b chl-a carotenoids chl-b phycocyanin phycoerythrin

Fig. 3.1. Wavelength spectrum showing the absorption spectra of chromoproteins present in the red algae; chl-a = chlorophyll a; chl-b = chlorophyll b (modified after Weiler & Nover 2008).

3.4 Phylogeny and evolutionary relationships of the Rhodophyta

The red algae are a very distinct group of organisms whose relationships to other groups are not certain, and the emergence of red algae may be the most ancient event in the evolution of eukaryotes (Hori & Osawa 1987). Possible evolutionary links between the red algae and the cyanobacteria (blue-green algae) have been suggested for several reasons. Both groups have phycoerythrin and phycocyanin as accessory pigments, both groups lack flagella, and both have non-aggregated thylakoids (Gabrielson et al. 1985). However, the cyanobacteria are prokaryotic (and thus lack chloroplasts) and they store food reserves as cyanophycean starch, an α1-4, α1-6 linked glucan resembling glycogen (Woelkerling 1990). The first occurrence of the red algae still remains unclear; Tappan (1980) has suggested that the red algae probably have existed for 2 billion years, while Hori & Osawa (1987) have estimated 1.3 to 1.4 billion years. The fossil record of the red algae is extremely uneven and only a small minority of taxa are represented (Tappan 1980). Herein, the most commonly recorded red algal fossils are taxa of the order Corallinales, which appear in the Early Cretaceous (Moussavian et al. 1993; Arias et al. 1995; Aguirre et al. 2000) and include more than 60 exclusively fossil genera, as well as at least 13 living genera to which fossil species have been assigned (Wray 1977; Johansen 1981; Woelkerling 1988).

12 Introduction to red algae (Rhodophyta)

However, in an analysis of the 57 exclusively fossil non-geniculate genera, Woelkerling (1988) has concluded that virtually all were of doubtful taxonomic status. Most other groups of the fossil red algae are attended by the same problems, thus making detailed evolutionary interpretations difficult (Woelkerling 1990). Despite those problems, the importance of the Corallinaceae is obvious and since all red algal species examined during the present study belong to this order, their characteristics are adressed below.

3.5 Geological record and evolution of the Corallinales

The order of the Corallinales belongs to the division of the Rhodophyta and appears in the Early Cretaceous (Moussavian et al. 1993; Arias et al. 1995; Aguirre et al. 2000). Other records of so-called “ancestral Corallinales” are reported e.g. from the Permian (Wu 1991) and the Triassic (Senowbari-Daryan & Velledits 2007) but are of doubtful taxonomic status, so this study will follow the findings of Aguirre et al. 2000. The Corallinales is a monophyletic group consisting of two families, the Corallinacea Lamou- roux, 1812 and the Sporolithaceae (Verheij, 1993). As stated above, examples of coralline- like algae older than the Early Cretaceous remain doubtful, and the earliest confirmed Meso- zoic record is Sporolithon rude (Lemoine) Ghosh & Maithy, 1996 from the Hauterivian (Arias et al. 1995), which belongs to the Sporolithaceae. Sporolithaceae and Corallinaceae are indistinguishable regarding vegetative anatomy but differ in the sporangial structure and spore division (Townsend et al. 1995). Due to the occurence of Sporolithon rude in the Early Cretaceous and its overall similarity to the Corallinaceae, it is likely that the Sporolitaceae are the ancestral group of the Corallinaceae (Aguirre et al. 2000). The Corallinales originated in the western tropical Tethyan realm, and the South Atlantic opening during the late Early Cretaceous favoured their geographic expansion, so they rapidly colonised the western and central Tethys and the western coast of central Africa during the Albian (Romanes 1916; Aguirre et al. 2000). The distribution records show that the Corallinales did not remain restricted to shallow-water reef ecosystems but also colonised deeper areas of carbonate platforms by the Albian (Agu- irre et al. 2000). Thus, the Corallinales seem to have followed a macroevolutionary onshore- offshore trend similar to that postulated for marine invertebrate orders (Jablonski & Bot- tjer 1990; Aguirre et al. 2000). The major species diversification history of the Corallinales extended from the latest Creta- ceous to the earliest Miocene, while the mid-Miocene to Pleistocene was a period of diversity stasis or even a slight decrease (Aguirre et al. 2000). During the Early Cretaceous, the Sporo- lithaceae and the Corallinaceae were equally diverse and during the Cenomanian temperature increase, the Sporolithaceae diversified further to their acme in the Coniacean. During the Maastrichtian and Daanian cooling, they were replaced as the most species rich group by a coralline subfamily, the cool- and deep-water Melobesioideae Bizzozero, 1885 (Agu- irre et al. 2000). From the Oligocene to the Early Miocene, the diversity of two other coral-

13 Introduction to red algae (Rhodophyta) line subfamilies, the Lithophylloideae Setchell, 1943 and the Mastophoroideae Setchell, 1943, rapidly increased (Aguirre et al. 2000), likely reflecting the partitioning of shallow- water habitats due to the onset of Southern Hemisphere glaciation near the Eocene-Oligo- cene boundary (Zachos et al. 1996). Coequally, the Sporolithaceae sharply decreased and the Melobesioideae further increased in species diversity in the Early Oligocene (Aguirre et al. 2000). After little change during the Miocene-Pleistocene, the onset of the Northern Hemi- sphere glaciation resulted in a decline of the Melobesioideae and a recovery of the Sporo- lithaceae. Parallel, the diversity of the Lithophylloideae and the Mastophoroideae increased to a Pleistocene maximum (Shackleton et al. 1984; Aguirre et al. 2000). At the present time, the Melobesioideae dominate the high-latitude ecosystems while the Mastophoroideae and Lithophylloideae are common in the warm-temperate and tropical areas, where the Melobesioideae and even the Sporolithaceae are reduced to greater depths and cryptic habitats (Fig. 3.2; Aguirre et al. 2000). The summary of coralline species diversity from the Cretaceous to the Pleistocene is shown in Fig. 3.3, clearly illustrating that the coralline algae still are major components of benthic marine communities. This is not least due to their adaptations to variuos environmental controls, particularly their withstands against the grazing pressure by many herbivores, which are highlighted in the following section.

polar temperate tropical temperate polar cold warm warm cold

Mastophoroideae Melobesioideae + Melobesioideae Lithophylloideae depth Sporolithaceae + Melobesioideae

Fig. 3.2. The principal present-day distribution of coralline algal subgroupings in dependence of water depth and geographical latitude (modified afterAguirre et al. 2000). Note that there is a considerable overlap between the subgroupings that is not shown in the figure.

14 Introduction to red algae (Rhodophyta)

0 Pleistocene lithophy./mastoph. Pl L Pl acme ELMELELM Za Me 10 To Se La

Miocene Bu 20 Aq

Ch melobesioid acme 30 Ru Oligocene

40 / Mastophoroideae Lithophylloideae

Lu Eocene 50 lithophylloid / mastophoroid expansion / mastophoroid lithophylloid

ELE Yp

60 Melobesioideae Da Paleocene

Ma 70 melobesioid expansion Time [Ma] Time Ca 80 Late Sa Co sporolith. 90 Sporolithaceae acme Tu

100 sporolithacean expansion sporolithacean Al

110 Cretaceous

Ap 120

Early Ba

130 Ha

140 Be ?

Fig. 3.3. Summary of rarefied coralline algal species diversity from the Cretaceous to the Pleistocene modified after Aguirre et al. 2000 with first confirmed records of Sporolithaceae and Corallinaceae being the Hauterivian and the Barremian respectively. A general long-term increase in coralline red algal species diversity from the Early Cretaceous to the Early Miocene is followed by a slight decline to the Pleistocene. Sporolithaceae dominate the Late Cretaceous (Coniacean acme) and Corallinaceae the Cenozoic (Aquitanian acme). Melobesioideae are the dominant subfamily during the Paleogene (Chattian acme) and are matched in species richness by Lithophylloideae and Mastophoroideae during most of the Neogene (Pleistocene acme).

15 Introduction to red algae (Rhodophyta)

3.6 Environmental adaptations of the Corallinales

The most successful order of the red algae are the heavily calcified coralline algae (Corallina- les Silva & Johansen, 1986), which dominate benthic marine communities under conditions where they (a) adhere to stable hard-substrata in shallow zones that have an abundance of herbivores; (b) adhere to stable hard-substrata in deep zones with few, if any, herbivores; and (c) occur at shallow and deep zones as free-living plants that are periodically overturned by wave action or currents, frequently in zones with reduced herbivory (Steneck 1986). Most of these coralline algal crusts are susceptible to fouling by fast growing fleshy algae and invertebrates in shallow productive zones (Littler 1972; 1976; Adey & Macintyre 1973; Jo- hansen 1981; Sebens 1986a; b), so grazing by herbivores is very often identified as a source of disturbance that keeps coralline algae clean and healthy (Doty 1959; Van den Hoek 1969; Paine & Vadas 1969; Littler 1972; Adey 1973; Steneck 1977; 1982; Paine 1980; Littler & Littler 1984; Padilla 1985). The positive correlation of herbivores with communities dominated by coralline algal crusts or rhodoliths is amply documented for the tropics (Littler 1972; Vine 1974; Lawrence 1975; Adey & Vassar 1975; Wanders 1977), temperate and boreal regions (Lawrence 1975; Townsend 1976; Raffaelli 1979; Paine 1980; Clokie & Boney 1980), and subarctic and arctic regions (Adey 1965; Steneck 1978; 1982; Brock 1979; Hagen 1983; Logan et al. 1984; Sebens 1986b). Since the coralline algae are not known to possess any secondary plant substances to prevent fouling by epiphytic organisms if herbivores are absent, they need other sources of physical disturbance, like abrasion by transport through currents (Steneck 1983). However, even if the mentioned disturbances are important for the coralline algae, they still cause stress for the organism itself. The three major herbivore groups feeding on coralline algae are molluscs, urchins, and fish, while the extent of herbivory exerted by these groups increased dramatically since the mid-Mesozoic Era (Steneck 1983). This intensification of grazing was part of the Mesozoic Marine Revolution (Vermeij 1977) that affected many groups of organisms and led to a kind of “arms race” between predators and prey. The three subfamilies of the coralline algae, being the Melobesioideae, the Mastophoroideae, and the Lithophylloideae, indicate a convergent evolution to withstand grazing pressure and physical disturbance, showing several anatomical and morphological characteristics (Table 3.3) that have been identified as contributing to their ecological success (Steneck 1986). Secondary pit connections and cell fusions (Fig. 3.4) that act as conduits for translocation are one important attribute required for the support of living cells beneath photosynthetic tissue, lateral growth, and recovery from deep wounds (Steneck 1983). In the group of the coralline algae, the white growing margins of a coralline crust and living cells more than 500 µm below the thallus surface are nonphotosynthetic and thus require translocation to remain metaboli- cally active, so thick and massively branched crusts could not survive without translocation (Steneck 1983). If coralline algal crusts are partially buried by sediment (e.g. maërl, Adey & Macintyre 1973; Bosence 1985) or overgrown by ephemeral organisms (Sebens 1986a), they also probably require translocation to support the stressed parts of the thallus. The recovery from deep wounds that remove outer photosynthetic tissue is also thought to be facilitated

16 Introduction to red algae (Rhodophyta)

Table 3.3. Convergently evolved adaptive characteristics and features of common coralline algal genera*; (X) reflects questionable placement in category (modified after Steneck 1986).

Morphology Conceptacles Multi-layered Adherence epithallium

Coralline taxa Thin Thick Branched Sunken Raised Yes No 100% 0% Melobesioideae Bizzozero, 1885 Lithothamnion Heydrich, 1897 XXXXXX XX Mesophyllum Lemoine, 1928 XXXXX X Phymatolithon Foslie, 1898 XXXXXX X Sporolithon Heydrich, 1897 XXXXXX X Lithophylloideae Setchell, 1943 Lithophyllum Philippi, 1837 XXXXXX XX Titanoderma Nägeli in Nägeli & Cramer, 1858 XXX X Mastophoroideae Setchell, 1943 Hydrolithon Foslie, 1909 XXXXXX XX

Neogoniolithon Setchell & Mason, 1943 X X X X X (X) XX Spongites Kützing, 1841a

Porolithon Foslie, 1909 X X X X X (X) XX Pseudolithophyllum Lemoine, 1913 X X (X) X X X X * Those having more than 10 species per genus, excluding epiphytes and parasites. a The genera of Neogoniolithon and Spongites were seperated (Woelkerling 1985) so the individual species are not all known.

by translocation (Steneck 1983). Another adaptation of many coralline algae is the so-called sunken meristem, which thus is protected against low-impact disturbance. The epithallium is the surface layer of cells that overlies and protects the intercalary meristem from which the epithallium and perithallium are derived (Fig. 3.4; Steneck 1982; 1985). The epithallia range in thickness from one (Adey & Macintyre 1973) to ten layers (Steneck 1986). If the grazing pressure mainly comes from limpets or chitons, thick coralline crusts with a deeply sunken meristem (multi-layered epithallium) dominate (Steneck 1985; Steneck & Paine 1986). Comparing the average bite-depths created by chitons (9 µm) and limpets (14 µm) (Steneck 1983) with the average epithallial thickness of associated coralline alge [e.g. up to 83 µm thick for Clathromorphum Foslie, 1898 (Adey 1965) and 51 µm for Lithophyllum Philippi, 1837 (Steneck & Paine 1986)] indicates that bites from these organisms are not able to penetrate the meristems of corallines that have a multi-layered epithallium. Thus, a multi-layered epithallium is apparently a very effective defence against high frequency but low intensity grazing such as that exerted by limpets and chitons (Steneck 1986). Further adaptations regard the morphology of the whole coralline algal thallus. In many non- tropical regions, stoutly branched crustose coralline algae predominate where the most intensive grazing is exerted by sea urchins (Townsend 1976; Milliken & Steneck 1981; Steneck 1982; 1985). Branched thalli are poor substrata for both adherence of and grazing by limpets and chitons (Milliken & Steneck 1981; Steneck 1982) and even limit the grazing pressure by sea urchins (Steneck 1986). In the North Atlantic, where sea urchin grazing is intense, an examination of grazing marks on branched thalli revealed that only 15% of the plant surface

17 Introduction to red algae (Rhodophyta)

(only the branch tips) was grazed

(Milliken & Steneck 1981). The epithallus thalli of branched rhodoliths and nodules may be protected from meristem intermediate levels of wave action in in a similar way than from herbivory (Littler & Littler 1984; perithallus Bosence 1985). The position of the conceptacles relative to the thallus surface is an- conceptacle other adaptation by the coralline algae against the grazing pressure exertet by herbivores. All coralline algal genera excluding one (Sporo- lithon Heydrich, 1897) produce spores and gametes within these conceptacles (Steneck 1986). At the time when spores and gametes are released, the conceptacle profile can be raised above, perithallus flush within, or sunken below the thallus surface (Fig. 3.4). The conceptacles of most coralline algae with thin and branched crusts hypothallus are raised, whereas those of many corallines with thick crusts are sunken below the thallus surface and hence are better protected fusion cells pit connections against the damage caused by grazing organisms, e.g. by the ra- Fig. 3.4. Schematic anatomy of the coralline algal thallus dulae of several chitons (Steneck with sunken conceptacles (modified after Steneck 1983); 1985; 1986). In non-tropical re- scale bars = 100 µm. gions with zones of intensive grazing pressure, raised conceptacles are only found on strongly branched crusts that hence are defended from such herbivore- induced disturbances (Steneck 1986). On the other hand, sunken conceptacles may not only be of advantage and it is likely that spore release is more efficient with raised conceptacles, and thus there seems to be a ‘reproductive cost’ for possessing sunken conceptacles (Steneck 1982). Hence, the feature of sunken conceptacles is rare e.g. in the tropics, and this also holds for thick, heavily grazed coralline algal crusts. The reason is that the prevailing parrotfish graze much deeper into the coralline thalli than even the most deeply sunken conceptacles, so there was no adaptive advantage but rather a disadvantage for sunken conceptacles (Adey

18 Introduction to red algae (Rhodophyta)

1965; Steneck 1983). Altogether, the heavily calcifying thallus is the most important attribute making the coralline algae resistant against abrasion by organisms or transport through currents (Steneck 1983). Though only the younger parts of the corallines bear living tissue while the older parts have already died back, the dead parts still outlast and with time, more and more calcified material becomes accumulated (Bosence 1983b). Because of that, coralline algae can grow very old and if the environmental conditions are suitable, they can form so-called rhodoliths (sensu Foster 2001), which are in the focus of the following chapter.

19 Introduction to rhodoliths

4. Introduction to rhodoliths

4.1 Definition, shape, and morphology of rhodoliths

Rhodoliths (Fig. 4.1) are free-living structures composed mostly (>50%) of non-geniculate coralline red algae that lack uncalcified joints (Adey & Macintyre 1973; Bosence 1983a; b). They are widely distributed in marine habitats and have an excellent fossil record since the Early Cre- taceous (Aguirre et al. 2000; Fos- ter 2001). One has to distinguish if a rhodolith is monospecific or multispecific, i.e. if it consists of one or more than one coralline algal species. Since they are intensely affected by currents, rhodoliths tend to grow in spheroidal, discoidal, or ellipsoidal shapes, thus resembling the hy- Fig. 4.1. Monospecific rhodoliths consisting of Lithotham- nion glaciale Kjellman from the Nordkappbukta site at drodynamic regime that predomi- 45 m water depth, where spheroidal shapes predominate nated during the growth process of (after Teichert et al. 2012); scale bar = 5 cm. the rhodoliths. This holds only for non-nucleated rhodoliths, while the shape of a nucleated rhodolith is widely determined by the nucleus, being e.g. a lithoclast (Freiwald 1995). The name rhodolith literally means “red stone” (Bosence 1983a) and consists of the ancient Greek words for rose-like (ῥόδειος) and stone (λίθοϚ). Structures composed of coralline algae are also known as maërl and mainly occur along the Atlantic coasts of Ireland and Scotland (Blunden et al. 1981). Maërl is a Breton term for unattached coralline algae and red algal gravels found off the northwestern coast of France (Bosence 1983b). Rhodoliths can develop from broken-off fragments of e.g. rigid coralline algal buildups (Frei- wald 1993; 1995) or from spore settlement on various solid materials (Bosence 1983a). Both kinds lead to different developments of the particular rhodoliths, and are explained in the next section. If the algal spores start to settle on a substratum, they are considered as an initial stadium of coralline algal crusts (Bosence 1991). During that state, they show some similari- ties to other coated grains like oncoliths (blue green algal nodules), and despite the clear dif- ference between coralline algal crusts and oncoliths, some authors have confused these terms (Bosence 1983a).

20 Introduction to rhodoliths

Rhodoliths are slow growing and can be long lived (>100 years), being resilient to varying environmental disturbances (Bosence 1983a; b). If they occur at high densities, these free- living non-geniculate coralline algae form rhodolith beds (Nelson et al. 2009), which are communities of high diversity due to their function as autogenic ecosystem engineers sensu Jones et al. (1994), meaning that they create habitats by their own skeletal growth and the successive modification of this skeleton After measuring the rhodoliths long (L), short (S), and intermediate (I) axes and applying the maximum projection sphericity formula [(S2/LI)0.5], the shapes can be quantified by plotting the gained values into a pebble shape diagram (Sneed & Folk 1958; Bosence 1983a) in order to draw conclusions about the hydrodynamic regime. Furthermore, the size of a rhodolith can be measured using the volume of an ellipsoid [(LIS/4π)0.5; Bosence 1976]. The internal structure of a rhodolith is often complex, e.g. the core can be multispecific and the outer layers can be monospecific if growth conditions got unsuitable for one ore more of the involved coralline algal species. If the details of such a succession are recorded, they may help to reveal the changing environmental conditions during the history of a rhodolith (Bo- sence 1983a). Considering what has been said, it is clear that the key to a correct reconstruction of the environmental attributes influencing the growth of a rhodolith is the knowledge of their growth mechanism. Since coralline algae are heavily calcified organisms, their biomineralisation process plays an important role and is explained in the next section, together with the features affecting the growth processes of rhodoliths.

4.2 Biomineralisation and growth mechanisms of rhodoliths

The rhodolith forming coralline algae are the most consistently and heavily calcified group of the red algae, and as such have been elevated to ordinal status (Corallinales Silva & Johans- en, 1986). Their calcification process involves high magnesium calcite precipitation within the cell walls (Bosence 1991). According to Alexandersson (1974; 1977) and Okazaki et al. (1982), coralline algae induce a microenvironment suitable for carbonate precipitation by metabolic excretion of alginic acid, so that the calcite saturation of the ambient water column is of minor importance for the calcification process. Early electron microscope work showed that the calcified cell walls of coralline algae have a two-layered structure, with an inner layer of acicular calcite parallel to the cell wall, succeeded by radial, inward growing calcite crystals (Fig. 4.2; Bailey & Bisalputra 1970; Alexanders- son 1974; 1977; Flajs 1977a; b; Garbary 1978; Cabioch & Giraud 1986; Bosence 1991). Microprobe plotting of magnesium and calcium concentrations by Flajs (1977b) and Mas- sieux et al. (1983) indicated that the secondary perpendicular layer has a higher magnesium content than the earlier parallel layer. The matrix that regulates the biomineralisation process is composed of glycoproteins (Cabioch & Giraud 1986). Coralline algae can exhibit a distinct rhythmicity in their growth pattern, that is visible in longitudinal sections (Fig. 4.3; Freiwald 1993). Freiwald & Henrich (1994) point out that this

21 Introduction to rhodoliths

early needle crystals later needle crystals Epithallus annual cycle annual cycle Perithallus

Fig. 4.2. Sheme of coralline algal cell wall calcification in two steps with early needle calcite parallel to cell walls and later needle calcite perpendicular to cell walls; slightly modified after Bosence (1991); scale bar = 2 µm. pattern represents annual increments comprising one dark band formed during winter and one bright band formed during summer. The colour of the dark bands results from an increased production of glycoproteins that coincides with an increased nutrient amount in the water column during the winter time (Freiwald 1993). These areas of intensified glycoprotein-production can be stained with fuchsine solution, that fur- Fig. 4.3. Coralline algal branch (Li- ther amplifies the visible banding pattern (Freiwald thothamnion glaciale Kjellman, 1883) 1993). from northern Norway in longitudinal section. The annual increments are The bands themselfes consist of several cell rows, clearly visible and consist of one dark which differ in their calcification level. The increments winter band and one bright summer start with dense, heavily calcified cell rows, grading band each; changed after Freiwald into less calcified cell rows. Since biomineralisation (1993); scale bar = 1 mm. proceeds throughout the growth period, the earlier formed cell rows are further calcified than cell rows formed towards the end of the growth period. Because ongoing calcification coincides with the production of the secondary cell layer (Bosence 1991), which has a higher magnesium

22 Introduction to rhodoliths content than the earlier magnesium-poor layer (Flajs 1977b; Massieux et al. 1983), the Mg/Ca ratio results in a correlation of high Mg/Ca values with heavily calcified cell walls and low Mg/Ca values with less calcified cell walls. The coralline algal biomineralisation processes develop a very rigid skeleton being resistant against physical impacts (Bosence 1983a; b), and if the environmental conditions are suitable, the coralline thalli can further develop to rhodoliths (Bosence 1991). Rhodoliths can either grow from detached fragments of existing rhodoliths or contain a core consisting of other material, a so-called nucleus, if they originate from the recruitment of coralline algal spores on hard substrata (Freiwald 1995; Gagnon et al. 2012). In the case of e.g. Lithothamnion glaciale Kjellman, 1883, which is in the focus of this study, the thalli of the coralline algae develop to thin, smooth crusts. After that and under sufficient conditions, the crusts grow thicker and form hummocks, the so-called protuberances (Jackson 2003; Adey et al. 2005). With further development, this crust-stadium changes to the rhodolith-stadium, when the proportion of the coralline algal mass exceeds the 50% ratio in relation to the nucleus (Bosence 1983a). The growth process of a rhodolith, regardless if nucleated or non-nucleated, involves its composition, shape, and structure. According to Bosence (1983a), the composition can be mono- or multispecific, depending on the number of involved coralline algal species. The shape varies between spheroidal, ellipsoidal, and discoidal forms and has to be determined as described in the previous section. Finally, the strucure of a rhodolith can be laminar, branching, or columnar, and this feature also depends on the growth mechanisms of the involved coralline algal species (Bosence 1983a). All three attributes, composition, shape, and structure of a particular rhodolith can vary over space and time (Bosence 1983a), so free-living rhodoliths can be very sensitive indicators of turbulence in the marine environment and the morphology within any species can vary from open to densely branched forms (Bosence 1991).

rhodolith form radial branching concentric crusts

increasing energy and turning

Fig. 4.4. Increase of lateral branch growth and branching density of rhodoliths with increasing hydrodynamic energy and turning frequency; slightly modified after Bosence (1991).

23 Introduction to rhodoliths

Densely branched and laminar rhodoliths are the result of frequent turning and high energy environments, since the apical abrasion of branches during turning leads to an intercalary branching or lateral filament growth below the apex (Bosence 1983b). This results in the development of typically spheroidal rhodoliths (Fig. 4.4) and may be followed by a concentric crustose growth of the original algal thallus or other encrusting organisms (Bosence 1991). The mechanisms mentioned above hold for both, nucleated and non-nucleated rhodoliths, but the future shape of a nucleated rhodolith is widely predetermined by the shape of the nucleus, especially if it consists of lithoclastic material (Freiwald & Henrich 1994; Frei- wald 1995; Foster 2001). Spheroidal, non-rhodoliths must be interpreted as a mature stage of development (Freiwald 1995), and since coralline algae, especially at northern latitudes, grow very slow (<1 mm yr-1 per branch; Freiwald 1993), it must be realised that individual rhodoliths may have a long history (Bosence 1983a).

4.3 Geological, palaeontological, and climatological significance of rhodoliths

Geologists and palaeontologists are interested in rhodoliths because of their common record in calcareous sediments (Milliman 1977; Canals & Ballesteros 1997) and fossil deposits (Foster 2001). The latter have been of particular interest because rhodoliths perhaps were even more abundant in the geological past than today (Copper 1994), have an excellent fossil record since the Early Cretaceous (Aguirre et al. 2000), and hence often can be used to estimate palaeoenvironmental conditions (Foster 2001). At a coarse scale, the presence of fossil rhodolith communities may indicate in which photic zone the particular deposit was formed (Toomey 1985). Since many recent coralline algal genera have existed since the Miocene, Bosence (1991) suggests the following bathymetric ranges for particular genera that should be applicable for the Neogene:

• Intertidal - 20 m: Neogoniolithon, Porolithon, Lithophyllum, Hydrolithon • 20 - 40 m: Neogoniolithon, Lithophyllum, Hydrolithon, Titanoderma, Mesophyllum • 40 - 60 m: Mesophyllum, Archaeolithothamnium, Lithothamnion, Lithophyllum • 60 - 100 m: Mesophyllum, Lithothamnion, Archaeolithothamnium, Lithophyllum

At finer scales, the rhodoliths may indicate water depth, sea-level stand, and particular features of the hydrodynamic regime (Bosence 1983a; Manker & Carter 1987; Braga & Martin 1988; Macintyre et al. 1991; Iryu 1997). In this coherence, the shape of rhodoliths has been of particular interest because many observations indicate that it can be affected by water motion (see above), so e.g. spherical shapes occur where water motion, and thus the repeated turning of the rhodoliths, is high (Bosence 1983b). However, these relationships may vary since spherical shapes also occur in low-energy environments (Reid & Macintyre 1988; Prager & Ginsburg 1989; Littler et al. 1991). Movement and shape of the rhodo-

24 Introduction to rhodoliths liths can be affected by many variables (Marrack 1991), so interpretations of hydrodynamic regimes based on the shape of rhodoliths should always be made with care (Foster 2001). Especially if rhodoliths are nucleated, their shape is widely predetermined by the core material (Freiwald 1995). As stated by Adey & Macintyre (1973), the accuracy of palaeoecological interpretations will be enhanced as the biology and ecology of living rhodoliths are better understood. Regard- ing their potential as climate recorders, rhodoliths could become the “Giant Sequoias of the sea” (Foster 2001: 665). The available data suggest that rhodoliths commonly grow less than one millimetre per year and leave behind growth increments like trees and corals (Freiwald 1993; Foster 2001). Frantz et al. (2000) determined the growth rate of Lithothamnion crassiusculum (Foslie) Mason, 1943 to 0.6 mm yr-1, using 14C accelerator mass spectrometry at 1-mm increments along a longitudinal branch section from tip to core. This indicated an age for large individuals of L. crassiusculum of c. 125 years. Frantz et al. (2000) also found 14C declines coincident with recent El Niño events, indicating that non-geniculate coralline algae can contain records of marine shallow-water climate similar to those found in corals (Linsey et al. 2000). Large, old rhodoliths and their encrusting counterparts were also successfully used to provide palaeoclimatic records concerning climate variability (Halfar et al. 2007; Hetzinger et al. 2009).

4.4 Geographical distribution of recent rhodolith beds

Rhodolith beds are common and widely distributed, and new reports of living rhodolith beds occur regularly due to enhanced research methods (Fig. 4.5). Rhodolith beds are particularly abundant in the Mediterranean, the Gulf of California, southern Japan, western Australia, and along the Atlantic coasts of Norway, Ireland, Scotland, north-eastern Canada, eastern Caribbean, and Brazil (Foster 2001). The largest known beds occur on the Brazilian Shelf and stretch from 2°N to 25°S (Kempf 1970; Milliman 1977). Recent, broad-scale surveys at the west coast of Ireland using remote sensing indicate that rhodolith beds occupy nearly 60 * 106 m2 of sea floor (De Grave et al. 2000), suggesting that similar surveys elsewhere may greatly increase the estimated abundances (Foster 2001). High concentrations of living rhodoliths can be found from the low intertidal zone down to 150 m water depth, typically in areas where the light regime is suitable for growth. Water movement has to be high enough to inhibit burial by sediment but coequally may not be too high or unidirectional to cause me- chanical destruction or rapid transport out of the favourable growing conditions (Steller & Foster 1995; Foster 2001). The temperature is the primary determinant of the geographical distribution of the rhodolith forming coralline algal species and the boundaries of biogeographical regions are associated with isotherms (Lüning 1990). However, temperature clearly does not control the distribution of rhodoliths in general as they are abundant from the Arctic to the tropics. Furthermore, the coralline algal species composition preserved in rhodoliths can reflect the climatic conditions prevailing during the time of rhodolith formation (Foster 2001).

25 Introduction to rhodoliths

Fig. 4.5. The world distribution of living rhodolith beds. The yellow circles indicate published rhodolith beds according to the compilations in Bosence (1983b) and Foster (2001); the green squares indicate the rhodolith beds examined in this study.

26 Introduction to cool-water carbonates

5. Introduction to cool-water carbonates

5.1 A short history of cool-water carbonate research

The first synthesis about cool-water carbonates by Chave (1967) emphasised that carbonates could form at all geographical latitudes, regardless of water temperature, as long as the terrigenous clastic sediment input was low. Even if cool-water carbonates have always been part of sedimentary geology (Flügel 1982), they were largely ignored between the 1950s and 1980s (James 1997). The results of ongoing research in the 1980s were subsequently published by Nelson (1988) and henceforward, there has been a surge in the number of studies regarding cool-water carbonates (James 1997).

5.2 Definition of cool-water carbonates

Cool-water carbonate factories almost exclusively produce biotically-controlled precipitates and the dominating organisms are heterotrophic (Schlager 2005). Though, the proportion of carbonate produced by photoautotrophic organisms, e.g. by coralline algae (Freiwald 1993; Henrich et al. 1997) can be substantial. Cool-water carbonate sediments typically consist of skeletal material of sand- to pebble-size while carbonate mud and abiotic marine cements are rare, and shallow-water reefs and oolites are absent (Schlager 2005). Cool-water carbonate factories extend poleward from the limit of tropical carbonate factories (c. 30° geographical latitude), whereat the transition between the cool-water and the tropical carbonate factories is gradual (Betzler et al. 1997; Schlager 2005). If cool-water carbonate factories occur at lower latitudes, they reside below the ther- mocline of the warm surface waters and in upwelling areas (Schlager 2005). The light regime in cool-water carbonate factories can be photic or aphotic, and the water has to be cold enough to exclude potential competition with organisms from the tropical carbonate factories, and it has to be sufficiently winnowed to prevent burial by terrigenous sediments (Schlager 2005). Compared to the tropical carbonate factories, the nutrient levels are generally higher in the cool-water carbonate factories (James 1997; Schlager 2005).

5.3 Controls on cool-water carbonates

Modern cool-water carbonates accumulate in seawater that is generally colder than 20°C, being part of the heterozoan association sensu James (1997). They are produced by coralline algae and benthic invertebrates, which feed through a variety of heterotrophic means, and can ocur worldwide in all platform environments (James 1997). The heterozoan association dif- fers from the light-dependent photozoan association sensu James (1997), that reflects shallow,

27 Introduction to cool-water carbonates warm-water, benthic calcareous communities, and both associations include modern and ancient skeletal and non-skeletal grain assemblages or facies (Fig. 5.1; James 1997). Carbonate sediments can only accumulate where the terrigenous clastic sediment input is limited (Chave 1967), so the rate of carbonate production versus the rate of terrestrial clastic sediment input is critical (James 1997). On the other hand, Nelson (1988) points out that terrigenous sediment load also carries nutrients with it, so cool-water carbonate sedimentation is favoured in areas with reduced but still existing terrigenous input. Because oceanic gyres transport cold polar waters equatorwards along the eastern sides of the modern oceans, cool-water carbonates are more common at those regions. This is amplified by the upwelling at the eastern margins of the modern oceans, leading to a cooling of the shelf waters and the input of nutrients, favouring the dominance of heterozoan sediments (James 1997). The distribution of cool-water carbonates also depends on the light regime. The lower boundary of the euphotic zone (1% surface illumination) is deep in oligotrophic environments, shallower in mesotrophic environments and shallowest in eutrophic environments, because an increased content of nutrients and hence a increased biomass in the water column leads to a reduced light penetration (James 1997). The photoautotrophic main producers of the tropic carbonate factories, green algae and zooxanthellate corals, are confined to the euphotic zone (James 1997). Below, the heterozoan assemblage prevails, while a peculiarity is given by the photoautotrophic coralline algae, that can tolerate a wider range of light levels than any other group of photosynthetic plants (Kain & Norton 1990), so they can prevail under light conditions that exclude other photoautotrophic organisms like the green algae. Because of that, coralline algae are equally important carbonate producers of cool-water carbonates as organisms from the heterozoan assemblage (Freiwald 1993; Henrich et al. 1997).

Heterozoan Photozoan association association

Molechfor Rhodalgal Chloralgal Coralgal facies facies facies facies

Bryomol Foramol Oopeloid Rudist facies facies facies facies

Bryonoderm Fusilinalgal facies facies

Fig. 5.1. The nomenclature of the heterozoan and photozoan particle association and the different facies types that can form each association; slightly modified after James (1997).

28 Introduction to cool-water carbonates

5.4 Cool-water carbonates in the Arctic

The formation of polar shelf carbonate deposits is well known from the Arctic and shows various occurrences, e.g. as rhodolith pavements from Mosselbukta at northern Svalbard (Kjellman 1883; 1885), bryozoans associated with siliceous sponges on Arctic shelves (Van Wagoner et al. 1989) and seamounts (Henrich et al. 1992; Henrich et al. 1995), and filter- feeding benthic communities consisting of barnacles and scallops on the southwestern Sval- bard shelf (Andruleit et al. 1996). While the rhodolith pavements from Mosselbukta at northern Svalbard, which were first discovered by Kjellman (1883) are a central aspect of this study, two of the other polar carbonate factories will be highlighted in the following. The facies belts and communities of the Arctic Vesterisbanken Seamount in the central Greenland Sea (73°30’N, 09°15’W; Henrich et al. 1992; Henrich et al. 1995) feature modern cool-water siliceous carbonate deposits. Despite the predominantly oligotrophic conditions, the seamount is extensively covered with biogenic mats, sponge bryozoan-serpulid buildups, bryozoan thickets, and sponge-crinoid mounds, thus resembling the modern end member of the foramol facies (Henrich et al. 1992). The environment of the mount exhibits low temperatures close to 0°C and salinities of c. 34.5, recorded nearly constant over the entire water column, which, together with the observation that all biogenic constructions are formed in the aphotic zone, suggests that there is no depth-related biogenic zonation on this seamount (Henrich et al. 1992). The fossilisation potential of the Vesterisbanken carbonates is argu- able, but Henrich et al. (1992) proposes that the resulting sediment after diagenesis might be a siliceous limestone that contains cool-water adapted pelagic and benthic organisms. This assumption may serve as a non-tropical altenative for similarly composed carbonate deposits of the fossil record, that also show low accumulation rates and a low species diversity (Hen- rich et al. 1992). The bioclastic carbonate sediments on the southwestern Svalbard shelf (76°45’N, 14°00’E; An- druleit et al. 1996) show the turnover from an infaunal softbottom community, consisting of a bivalve assemblage, towards an epibenthic hard substrate community c. 2400 years ago. The modern epibenthic community consists of barnacles, bivalves, bryozoans, brachiopods, serpulids, and kelp, thus the skeletal carbonate producers represent a member of the barnamol facies (Andruleit et al. 1996). The turnover from an infaunal to an epibenthic community was caused by the deglaciation during the Holocene (Elverhøi et al. 1990), leading to an increase of the hydrodynamic regime that washed out the soft sediments (Andruleit et al. 1996). This shows the strong relationship between the composition of the communities that produce the cool-water carbonates and the sedimentary environment, which, in this case, is glacio-marine influenced (Andruleit et al. 1996). Altogether one can say that Arctic cool-water carbonates are generally dominated by heterozoan organisms (James 1997), while photoautotrophic coralline algae are a peculiarity (Freiwald 1993; Freiwald & Henrich 1994). On this basis of accumulated evidence, fossil cool-water carbonates can be interpreted with increasing success (James 1997).

29 Material and methods

6. Material and methods

6.1 Overview

Samples and data were obtained from 31th July to 17th August in the year 2006 during the MSM 02/03 expedition of RV Maria S. Merian (Fig. 6.1a; see Table 6.1 for a station and gear list, and Lherminier et al. 2006 for the cruise report). It has to be stressed that the author of the present study did not take part in the expedition. All data acquisition carried out on board of RV Maria S. Merian was carried out by the participants of the cruise (For a list of the participants, please refer to Lherminier et al. 2006: 3).

6.2 Exploration of the seafloor

Seabed mapping was carried out with a Kongsberg® EM 1002 multibeam echo sounder (see Table 6.1 for the start and end points of the multibeam surveys), operating at a nominal sonar frequency of 95 kHz and controlled with the software package SIS® (Seafloor Infor- mation System). The multibeam raw data were processed using the software packages Nep- tune® and Cfloor®. The Neptune® bathymetric post-processing software brought raw multibeam data through a data correction and cleaning process and was designed for graphical description of the raw data to identify problems. It also provided tools to correct or remove errors in navigation data, depth soundings, and sound speed profiles. The Cfloor® software was used for chart production starting with Neptune® output data from which digital ter- rain models were generated. The manned submersible JAGO (Fig. 6.1b) was used for the visual inspection and video docu- mentation of shallow-water slopes, which were preselected in order to the acquired multibeam data, and for sampling of the rhodolith communities with a hydraulic manipulator arm (see Table 6.1 for the start and end points of the JAGO dive tracks). The submersible was certified to a maximum operating depth of 400 m and accommodated two persons, the pilot and a scientific observer.

6.3 Sampling of the water column

For the measurements of physical and chemical parameters in the water column, a Sea Bird SBE® CTD mounted with a rosette of 24 water bottles (10 L capacity each) was employed (Fig. 6.1c). Altogether, nine CTD surveys were carried out (see Table 6.1) and the water samples for total alkalinity (TA) and dissolved inorganic carbon (DIC) were taken at the maximum water depth at each station. Additional near-bottom water samples were obtained by a 5 L Niskin bottle attached to the JAGO submersible. The temperature was measured in °C, the reference

30 Material and methods

Table 6.1. Station and gear list of the expedition of RV Maria S. Merian MSM 02/03 showing start and end points of the used gear (CTD = measurements of conductivity, temperature, and depth; MB = multibeam echosounder; DRG = dredge; JAGO = manned submersible; LS = light measurements; Lat. = geographical latitude; Long. = geographical longitude).

Start End

Station # Gear Location Date Time (UTC) Lat. [°N] Long. [°E] Depth [m] Time (UTC) Lat. [°N] Long. [°E] Depth [m]

616 CTD Floskjeret 7/31/2006 10:18 78º09.29‘ 13º50.05‘ 424 10:55 78º09.29‘ 13º50.05‘ 424

632 CTD Krossfjorden 8/1/2006 23:36 79°02.37‘ 10°46.99‘ 340 00:09 79°02.37‘ 10°46.99‘ 340

633-1 MB Krossfjorden 8/2/2006 00:45 79°02.65‘ 10°45.57‘ 341 01:11 79°05.80‘ 10°43.26‘ 42

633-2 MB Krossfjorden 8/2/2006 01:17 79°05.77‘ 10°44.31‘ 49 01:46 79°02.70‘ 10°50.48‘ 332

633-3 MB Krossfjorden 8/2/2006 02:01 79°02.96‘ 10°54.97‘ 350 02:30 79°05.79‘ 10°44.91‘ 46

633-4 MB Krossfjorden 8/2/2006 02:35 79°05.79‘ 10°45.88‘ 53 03:06 79°03.42‘ 10°59.14‘ 308

633-5 MB Krossfjorden 8/2/2006 03:18 79°04.22‘ 11°01.06‘ 184 03:46 79°05.86‘ 10°46.47‘ 46

633-6 MB Krossfjorden 8/2/2006 03:52 79°05.91‘ 10°47.86‘ 53 04:17 79°04.81‘ 11°03.29‘ 110

633-7 MB Krossfjorden 8/2/2006 04:26 79°05.30‘ 11°04.03‘ 94 04:54 79°06.03‘ 10°48.37‘ 43

633-8 MB Krossfjorden 8/2/2006 05:01 79°06.16‘ 10°50.65‘ n.d. 05:21 79°05.69‘ 11°03.93‘ 93

633-9 MB Krossfjorden 8/2/2006 05:28 79°06.08‘ 11°03.37‘ 115 06:05 79°06.42‘ 10°42.34‘ 36

633-10 MB Krossfjorden 8/2/2006 06:05 79°06.42‘ 10°42.34‘ 36 06:38 79°06.37‘ 11°03.68‘ 130

634 JAGO Krossfjorden 8/2/2006 08:19 79°04.50‘ 10°47.20‘ 261 10:49 79°04.99‘ 10°47.65‘ 140

636 DRG Krossfjorden 8/2/2006 14:17 79°05.37‘ 10°47.22‘ 99 15:05 79°05.69‘ 10°49.44‘ 100

637 DRG Krossfjorden 8/2/2006 15:34 79°05.94‘ 10°48.21‘ 51 16:11 79°05.84‘ 10°46.36‘ 49

638-1 MB Krossfjorden 8/2/2006 17:27 79°04.66‘ 10°43.61‘ 213 19:53 79°05.92‘ 10°52.50‘ 136

638-2 MB Krossfjorden 8/2/2006 20:28 79°05.11‘ 10°56.19‘ 127 21:35 79°04.23‘ 11°25.67‘ n.d.

639 MB Krossfjorden 8/2/2006 21:35 79°04.23‘ 11°25.67‘ n.d. 08:00 79°05.69‘ 10°46.84‘ 70

640 CTD Krossfjorden 8/2/2006 23:56 79°18.32‘ 11°36.69‘ 132 00:21 79°18.32‘ 11°36.69‘ 132

641 CTD Krossfjorden 8/3/2006 01:32 79°12.99‘ 11°44.48‘ 243 01:46 79°12.99‘ 11°44.48‘ 243

642 CTD Krossfjorden 8/3/2006 02:36 79°09.10‘ 11°46.73‘ 350 03:10 79°09.10‘ 11°46.73‘ 350

643 CTD Krossfjorden 8/3/2006 04:05 79°05.55‘ 11°34.30‘ 283 04:33 79°05.54‘ 11°34.30‘ 284

644 JAGO Krossfjorden 8/3/2006 06:39 79°05.70‘ 10°46.74‘ 71 09:29 79°05.95‘ 10°45.61‘ 41

651 MB Krossfjorden 8/4/2006 01:46 79°12.87‘ 11°44.37‘ 234 05:40 79°03.64‘ 11°14.86‘ 106

652 JAGO Krossfjorden 8/4/2006 06:56 79°03.52‘ 11°14.05‘ 99 09:29 79°04.35‘ 11°14.43‘ 47

654 LS Krossfjorden 8/4/2006 12:56 79°05.45‘ 10°48.20‘ 95 n.d. n.d. n.d. n.d.

669 CTD Mosselbukta 8/6/2006 03:51 79°55.33’ 15°35.17’ 156 04:08 79°55.33’ 15°35.16’ 156

670 MB Mosselbukta 8/6/2006 06:28 79°54.20‘ 15°44.91‘ 81 10:33 79°53.40‘ 15°45.88‘ 42

671 JAGO Mosselbukta 8/6/2006 11:40 79°54.64‘ 15°48.62‘ 44 14:09 79°54.51‘ 15°50.23‘ 27

672 LS Mosselbukta 8/6/2006 11:40 79°54.57‘ 15°49.72‘ 44 13:00 79°54.57‘ 15°49.72‘ 44

675 DRG Mosselbukta 8/6/2006 20:57 79°54.21‘ 15°46.94‘ 61 21:59 79°53.52‘ 15°44.89‘ 72

684 JAGO Mosselbukta 8/7/2006 16:43 79°53.72‘ 15°44.75‘ 80 19:24 79°53.54‘ 15°46.58‘ 44

685 LS Mosselbukta 8/7/2006 16:55 79°53.73‘ 15°44.62‘ 79 17:34 79°53.61‘ 15°45.65‘ 61

699 CTD Nordkappbukta 8/9/2006 06:59 80°35.00‘ 19°27.84‘ 144 07:15 80°35.03‘ 19°27.48‘ 144

700 MB Nordkappbukta 8/9/2006 07:59 80°31.62‘ 19°39.45‘ n.d. 11:34 80°31.00‘ 19°44.09‘ 41

701 JAGO Nordkappbukta 8/9/2006 12:08 80°31.99’ 19°50.62‘ 73 15:24 80°31.95‘ 19°51.30‘ 30

702 LS Nordkappbukta 8/9/2006 12:28 80°32.04‘ 19°50.74‘ 94 12:46 80°32.00‘ 19°50.74‘ 75

703 JAGO Nordkappbukta 8/9/2006 15:57 80°31.93‘ 19°50.68‘ 57 16:43 80°31.77‘ 19°48.42‘ 95

704-1 DRG Nordkappbukta 8/9/2006 17:32 80°31.24‘ 19°44.32‘ 47 17:40 80°31.27‘ 19°43.84‘ 46

704-2 DRG Nordkappbukta 8/9/2006 18:07 80°31.22‘ 19°44.27‘ 45 18:29 80°31.44‘ 19°43.02‘ 44

706 CTD Nordkappbukta 8/9/2006 20:05 80°32.01‘ 19°50.77‘ 71 20:22 80°32.02‘ 19°50.77‘ 81

707 MB Nordkappbukta 8/9/2006 20:39 80°31.76‘ 19°49.34‘ 87 21:16 80°32.23‘ 19°36.74‘ 84

708 MB Nordkappbukta 8/9/2006 22:14 80°33.89‘ 18°27.07‘ 121 22:48 80°34.36‘ 18°06.29‘ 134

710 MB Nordkappbukta 8/10/2006 01:54 80°31.72‘ 19°39.79‘ 69 07:55 80°31.83‘ 19°39.40‘ 41

711 JAGO Nordkappbukta 8/10/2006 08:56 80°31.80‘ 19°40.67‘ 72 11:54 80°31.86‘ 19°41.47‘ 30

712 LS Nordkappbukta 8/10/2006 09:02 80°31.81‘ 19°40.63‘ 68 09:22 80°31.77‘ 19°40.32‘ 69

713 MB Nordkappbukta 8/10/2006 12:36 80°32.09‘ 19°43.49‘ 112 16:16 80°31.76‘ 19°44.06‘ 60

714 JAGO Nordkappbukta 8/10/2006 17:02 80°32.19‘ 19°50.40‘ n.d. 18:56 80°31.51 19°52.28’ n.d.

756 MB Floskjeret 8/17/2006 08:17 78°18.30‘ 14°34.32‘ 127 08:58 78°18.86‘ 14°31.94‘ 54

757 JAGO Floskjeret 8/17/2006 10:30 78°18.64‘ 14°32.00‘ 63 12:34 78°18.74‘ 14°31.22‘ 40

31 Material and methods system for salinity was the practical salinity scale (PSS). The TA was determined by potentiometric titration (Gran 1952) using a Metrohm Basic Titrino 794® system. The DIC was analysed by means of a SEAL QUAATRO® autoanalyser, following the method of Stoll et al. (2001). The results of both methods were corrected by parallel measurements of certified reference standards (Dickson et al. 2003). The carbonate system parameters pH, carbon dioxide partial pressure (pCO2), calcite saturation (ΩCal) and aragonite saturation (ΩArg) were calculated from TA, DIC, temperature, and salinity.

6.4 Measurements of the light regime

A total of five light profiles of the water column were obtained with a LI-COR Spherical Quantum® Sensor (LI-193SA) in combination with a LI-COR data logger (LI-1400) and a 100-m-long cable (Fig. 6.1d; see Table 6.1). The wavelength spectrum (400-700 nm) cor- responded to the photosynthetic active radiation (PAR; unit = µmol photons m-2 s-1). The mea-surements were undertaken from the small rescue boat of RV Maria S. Merian in order to minimise potential bias due to shadows cast by the research vessel and to enable controlled measurements just below the water surface. In addition to the subaquatic measurements, the surface PAR was measured and used as a reference for various weather conditions. The raw data were translated to percentages with respect to the surface illumination in order to determine the water depth of the photic zone boundaries at each study site.

6.5 Sampling of the benthic community

A dredge with an opening of 100 cm width and 40 cm height and a net of 0.5 cm mesh size was used to sample macro- and megabenthic epifauna (Fig. 6.1e). The samples obtained by the five dredgings (see Table 6.1) were sieved (1 mm), the organisms were collected from the overall catch and fixed in 70% alcohol for later analysis in the laboratory using a Zeiss Stemi® 2000 stereo microscope and a Leica Apozoom® 1:6 M420 macroscope with a Leica DFC-320 Digicam. The specimens were determined to the lowest possible taxonomi- cal level each.

6.6 Sampling of the rhodoliths

The rhodolith samples were collected by the hydraulic manipulator arm of the manned submersible JAGO (Fig. 6.1f) during the ten seafloor surveys (see Table 6.1 for the start and end points of the JAGO dive tracks) and the dredge (see previous section). The collected rhodoliths were assorted and only monospecific rhodoliths composed of Lithothamnion glaciale Kjellman with sufficient protuberance development were included to the calculation of the

32 Material and methods annual carbonate production. The samples were collected from one site at Floskjeret (757), three sites at Krossfjorden (637, 644, 652), one site at Mosselbukta (684), and three sites at Nordkappbukta (701, 711, 714). One sample from Nordkappbukta (714) was collected in significantly shallower water depth in order to check the influence of water depth on the annual carbonate production.

a b c

d e f

Fig. 6.1. Employed gear during the MSM 02/03 expedition of RV Maria S. Merian; (a) research vessel RV Maria S. Merian; (b) manned submersible JAGO; (c) CTD with mounted water bottles; (d) light measurements; (e) dredge; (f) rhodolith sampling with the JAGO hydraulic manipulator arm; images provided by the participants of the MSM 02/03 cruise.

6.7 Taxonomy of the coralline algae

The coralline algal genera and species were determined using thin sections (Zeiss Stemi® 2000 stereo microscope, Leica Apozoom® 1:6 M420 macroscope with Leica DFC-320 Digicam), histological slices (Zeiss Axiophot®), and samples for the scanning electron microscope (SEM, Tescan Vega® \\ XMU). For the preparation of the SEM samples, longitudinal broken protuberances were attached to aluminium stubs using PONAL express wood glue and were cleaned with compressed air after hardening. The samples were sputter coated with gold for 4 min using a Cressington 108auto sputter coater and were exam-

33 Material and methods ined with an electron beam energy of 20-30 kV. The identification of the coralline algal species was considerably accompanied and improved by Dr. William Woelkerling at La Trobe University, Department of Botany, Bundoora (Vic- toria), Australia. The identification of thalli of Lithothamnion glaciale Kjellman, 1883 was based on data in Adey (1964; 1966; 1970b, including species keys), Adey et al. (2005, including comparisons of L. glaciale with Lithothamnion tophiforme (Esper) Unger, 1858 and Lithothamnion lemoineae Adey, 1970), Irvine & Chamberlain (1994, including species keys and accounts), and Kjellman (1883; 1885, containing the original account of L. glaciale). The identification of specimens of Phymatolithon tenue (Rosenvinge) Düwel & Wegeberg, 1996 was based on Rosenvinge (1893) and Düwel & Wegeberg (1996). The JAGO video footage also showed the occurrence of non-calcareous red algae that were mostly finely filamentous and resembled Polysiphonia Greville, 1823 (Rhodomelaceae, Cer- amiales; see also Womersley 1979). As unfortunately, no specimens were collected during the 2006 cruise, unequivocal identification of these algae was not possible, and they are referred to hereafter as ‘Polysiphonia-like red algae’.

6.8 Description and classification of the rhodoliths

For the description and classification of the rhodoliths, specimens from Floskjeret (45 m water depth, n = 37), Krossfjorden (41-50 m water depth, n = 122), Mosselbukta (44 m water depth, n = 92), and Nordkappbukta ( 27-50 m water depth, n = 128) were used. The shape of the rhodoliths was analysed by measuring the long (L), intermediate (I) and short (S) axes of the specimens and applying the maximum projection sphericity formula [(S2/LI)0.5] as described in Bosence (1983a), and the results were plotted into a pebble shape diagram (Sneed & Folk 1958) using the MICROSOFT EXCEL® sheet Tri-plot (Graham & Midgley 2000). The size of the rhodoliths was measured using the volume of an ellipsoid [(LIS/4π)0.5] as described in Bosence (1976).

6.9 Computer tomography scans of the rhodoliths

The computer tomography scans were carried out at the Waldkrankenhaus in Erlangen, Ger- many with a SIEMENS SOMATOM® Sensation 16. Eight rhodoliths collected at the Mos- selbukta site in 44 m water depth were scanned with the non-destructive technique of computer tomography in order to reveal the organisms present inside hollow rhodoliths, attached to the rhodoliths surface, or overgrown by the coralline red algal carbonate skeleton. The rhodoliths were separately introduced into the CT in order to gain X-ray image slices reconstructed from the spiral CT-data set. The data acquisition was carried out with 90 mA at 120 kV and 2.0 mm slice width with a collimation of 16 x 0.75 mm. The data were saved in the conventional DICOM-format and the image reconstructions were carried out on a SIEMENS SYNGO® InSpace workstation. Two perpendicular longitudinal stacks of X-ray

34 Material and methods slice images (0.75 mm spacing) were reconstructed for each rhodolith and transverse slices were reconstructed likewise at 0.75 mm spacing. The reconstructions were carried out with the rather soft B20s (for abdomen) and the rather hard B70s (for bones) algorithms. X-ray slices were reconstructed for all scanned rhodoliths while ideal images were achieved with the settings trapezoidal ramp 73/51, pyramid 44/35, brightness 73, opacity 51, width 1243, and centre 2373. The data files were processed for later examination independent from the SIEMENS SYNGO® workstation using the portable software SIEMENS SYNGO® fastView.

6.10 Verification of the annual rhodolith banding pattern

Early electron microscope work showed that the calcified cell walls of coralline algae have a two- layered structure with an inner layer of acicular calcite parallel to the cell wall, succeeded by radial, inward growing calcite crystals (Bailey & Bisalputra 1970; Alexandersson 1974; 1977; Flajs 1977a; b; Garbary 1978; Cabioch & Giraud 1986; Bosence 1991). Microprobe plotting of magnesium and calcium concentrations by Flajs (1977b) and Massieux et al. (1983) indicated that the secondary radial layer has a higher magnesium content than the earlier parallel layer. Because of that, the Mg/Ca ratio should decrease towards the end of a growth period, where the cell rows are less calcified than at the beginning of the growth period. The growth increments from one protuberance of Lithothamnion glaciale were tested for their correlation with an annual pattern by measuring the Mg/Ca variation along the growth axis of rhodolith protuberances using electron dispersive X-ray (EDX) spectroscopy. The measurements were done with an Oxford Instruments X-Max® EDX with a 50 mm2 detector area mounted on a Tescan Vega® \\ XMU SEM with copper for specimen calibration. For the preparation of the SEM examination, a protuberance of L. glaciale was resin embedded, longitudinal cut using a water-cooled low speed diamond saw and wet polished with SiC powder (220, 400, and 800 graining). The specimen was attached to an aluminium stub using PONAL express wood glue and was cleaned with compressed air after hardening. The sample was sputter-coated with gold for 4 min using a Cressington 108auto sputter coater and was examined with an electron beam energy of 20 kV, while spots for Mg/Ca measurements were selected at 15 µm intervals. The EDX measurements were carried out for a time interval of 120 s per selected spot. Ad- ditionally, the annual forming, spore-bearing conceptacles (Jackson 2003) were checked for their parallel succession relative to the growth increments.

6.11 Calculation of the annual carbonate production by the rhodolith beds

First, the collected rhodoliths from all sites were sorted and checked for their sufficiency, i.e. that they were monospecific and consisted of Lithothamnion glaciale. The living surfaces of the sufficient rhodoliths (Floskjeret, n = 17; Krossfjorden, n = 48; Mosselbukta, n = 27; Nordkappbukta, n = 75) together with a 5 cm scale were photographed using a Zeiss 50

35 Material and methods mm Macro mounted on a Nikon D200 DSLR and the software Nikon Capture Con- trol®. The software Olympus analySIS FIVE® was used to calibrate the images, to measure the area of the living surface, and to count the protuberances (being the main spots of rhodolith CaCO3 production) on the living surface. This approach resulted in the median numbers of protuberances per area [n m-2], which were checked for coherence using Levene’s test for homogeneity of variance based on means (One-way ANOVA). Five protuberances each of randomly selected rhodoliths were cut off, longitudinal measured using a Hazet digital calliper, dried in a Memmert 700 cabinet desiccator at 40°C for 48 h, and weighed using a Mettler Toledo® AB204-S classic precision balance. The processed protuberances were resin embedded, longitudinal cut using a water-cooled low speed diamond saw, and wet polished with SiC powder (220, 400, and 800 graining). The polished sections were etched with 0.1 n HCl for 3 s and stained with fuchsine solution (1 g fuchsine in 100 mL ethanol 50%) for 10 s to amplify the dark areas of increased glycoprotein-production during the winter time and therefore to visualise the annual growth increments. The growth increments per protuberance were counted using a Zeiss Stemi® 2000 stereo microscope. If growth increments could not be distinguished clearly, the annual forming conceptacles (Jackson 2003) were used as a substitute. The counted increments were plotted against the weight of the whole protuberance to check if the growth succession was linear using the reduced major axis algorithm and Levene’s test for homogeneity of variance based on means (One-way ANOVA).

Subsequent band counting led to the mean amount of produced CaCO3 per protuberance -1 per year [g CaCO3 yr ] which was checked for significance using Levene’s test for homogeneity of variance based on means (One-way ANOVA). Together with the calculated number of protuberances per year, the annual carbonate production per square metre per year -2 -1 [g CaCO3 m yr ] was calculated by multiplication of the values. The results were plotted against the parameters water depth, geographical latitude, duration of the polar night, duration of sea ice cover, annual mean temperature, and seawater calcite saturation of the respec- tive sampling site and checked for coherence using the reduced major axis algorithm and Levene’s test for homogeneity of variance based on means (One-way ANOVA). All statistical analyses were carried out using the palaeontological statistics software package PAST (Ham- mer & Harper 2001).

36 Results

7. Results

7.1 Seafloor topography and distribution of rhodolith beds

The Floskjeret site (78°18’N, 14°32’E) at the mouth of Borebukta mapped by multibeam echosounder covers c. 0.4 km2. The seafloor rises constantly from SE to NW and the variably sized depressions are presumably iceberg grounding-marks (Fig. 7.1). Dive track 757 (60-40 m water depth) heads towards the inner part of Borebukta and shows a smooth seafloor covered with presumably glaciogenic lithoclasts of cobble size and bio- clasts mainly consisting of shells, barnacles and sea urchin tests, covered with fine sediment. Boulders (>1 m3) occur occasionally and are the only prominent morphological character on the otherwise flat ground. In 60 m water depth, the coverage with thin coralline algal crusts

757 30 depth [m] 78°18.4‘ 78°18.6‘ 78°18.8‘

= JAGO dive track

757 = Station number 200 m

130 N 14°32‘ 14°33‘ E

Fig 7.1. Multibeam map of the Floskjeret site showing depth and morphology of the seafloor and the positions of the applied gear (see Table 6.1 for a list of the kind and position of each applied gear); starting point of the track is the numbered symbol.

37 Results

(CAC) is rare (<5%) and limited to smooth crusts without protuberances (Fig 7.2a). This coverage increases with decreasing water depth to up to 100% in 47 m water depth, where CAC with well-developed protuberances (>5 mm length) and rhodoliths up to 20 cm in diameter occur (Fig 7.2b). Coequally, the coverage with fine sediments decreases. These conditions do not change up to 40 m water depth.

Fig. 7.2. JAGO-photographs of the seafloor at Floskjeret; (a) CAC without protuberances growing on cobbles in 60 m water depth; (b) CAC with well-developed protuberances growing on cobbles together with rhodoliths in 47 m water depth; scale bars = 10 cm.

38 Results

The Krossfjorden site (79°08’N, 11°40’E) mapped by multi- 79°20‘ 2 beam echosounder covers c. 176 km and consists of Lil- 647 liehöökfjorden, Krossfjorden and Mitragrunnen Bank at the mouth of Krossfjorden (Fig. 7.3). The inner part of Lilliehöök- fjorden begins with the steep glacier slope of Lilliehöökbreen 640 79°18‘ in c. 150 m water depth and leads into a narrow channel with smooth morphology and c. 270 m water depth. When the channel turns to the SW, it passes into Krossfjorden, and the morphology changes to a steep moraine ridge in c. 50 m wa- 79°16‘ ter depth at the entrance of the fjord. Beyond this ridge, the seafloor rapidly rises in direction to the even Mitragrunnen

Bank. Iceberg scouring-marks are visible over the entire site, Lilliehöök orden but mainly in water depths >100 m. 79°14‘ Dive track 634 (250-140 m water depth) south of Mitragrun- nen Bank shows presumably glaciogenic pebbles and cobbles 641 covered with fine sediment, while boulders (>1 3m ) occur only occasionally. The well-sorted lithoclastic pavement is 79°12‘ regularly grooved by sediment trapping iceberg scouring- marks, which are oriented parallel to the slope. The slopes inclination is c. 20° on average. With decreasing water depth, the amount of the fine sediment decreases, the coarse mate- 79°10‘ rial becomes less sorted, and the amount of shells increases. CAC and rhodoliths are absent. 642 79°08‘

= JAGO dive track Mitragrunnen

= Dredge track

= Light measurements Kross orden 637 654 = CTD measurements 79°06‘ 643 644 636 643 = Station number

30 634 79°04‘ 652 depth [m]

CTD - transect 632 2000 m 380 N 10°45‘ 11°00‘ 11°15‘ 11°30‘ 11°45‘ E

Fig 7.3. Multibeam map of the Krossfjorden site showing depth and morphology of the seafloor and the positions of the applied gear (see Table 6.1 for a list of the kind and position of each applied gear); starting point of a track is always the numbered symbol; the dashed line marks the CTD-transect referred to in the next chapter (see Fig. 7.12a; b).

39 Results

Along dive track 644 (60-41 m water depth), situated at the margin of Mitragrunnen Bank, patches of pebbles and cobbles alternate with depressions up to 1 m in depth and up to 15 m in diameter. These depressions contain shell accumulations, mainly from Chlamys islandica Müller, 1776 and Hiatella arctica Linnaeus, 1767 on fine sediment. CAC appear at 58 m water depth (c. 5% coverage) and c. 20% of the crusts have protuberances. CAC are restricted to pebbles and cobbles while fine sediment depressions are not colonised. From a water depth of 53 m upwards, CAC coverage on hard substrate increases to 25-75%, and up to 50% of the CAC have well-developed protuberances. Rhodoliths are absent. Dive track 647 (41-39 m water depth) at the glacier slope of Lilliehöökbreen shows a fine sediment seafloor with a distinct morphology made up by mounds and trenches. The sediment has a very fine grain size and is laced all over with burrows of unknown origin (Fig. 7.4). This fine sediment presumably origins directly from the melting glacier front, and the general morphology appears to change regularly as a result of the glacier dynamics. Iceberg scouring-marks occur very regularly but are poorly preserved due to the unstable nature of the sediment. Lithoclastic hard substrate like pebbles and cobbles, shell accumulations, CAC, and rhodoliths are not detectable. Dive track 652 (92-47 m water depth) heads along the moraine ridge at the entrance of Kross- fjorden and shows distinct iceberg scouring-marks perpendicular to the ridge. The seafloor features pebbles and cobbles while boulders (>1 m3) are rare. Accumulations of shells, mainly from C. islandica and H. arctica, and fine sediment is limited to small depressions (up to 0.5 m in depth and up to 5 m in diameter) and to the iceberg scouring-marks. CAC appear at 77 m water depth (<5% surface coverage; Fig. 7.5a) and constantly increase in coverage and in development of their protuberances. In 50 m water depth, c. 40-80% of the hard substrate is covered with CAC, generally with well-developed protuberances. Coequally, several rhodoliths up to 15 cm in diameter occur but are rare (Fig. 7.5b).

Fig. 7.4. JAGO-photograph of the seafloor at the glacier slope of Lilliehöökbreen in 40 m water depth, showing a fine sediment seafloor with burrows of unknown origin and ophiurids; scale bar = 10 cm.

40 Results a

Fig. 7.5. JAGO-photographs of the seafloor at Krossfjorden; (a) initial CAC without protuberances together with shells of C. islandica (living) and H. arctica (dead) and barnacles (living) in 77 m water depth; (b) well-developed CAC and rhodoliths together with shells of C. islandica (dead) and barnacles (living) in 50 m water depth; scale bars = 10 cm.

41 Results

The Mosselbukta site (79°53’N, 15°55’E) mapped by multibeam echosounder covers c. 13 km2. The seafloor continuously rises from c. 170 m to c. 20 m water depth towards the shore in the SE. It shows a strong morphology with distinct ridges heading parallel to the shoreline. Iceberg scouring-marks are clearly visible and head perpendicular to the shoreline (Fig. 7.6). Along dive track 671 (42-27 m water depth), perpendicular to the southern Mosselbukta shoreline, 80-100% of the seafloor at 42 m water depth are covered with coralline algae, partly in form of CAC, but mostly as rhodoliths (Fig. 7.7a). The rhodoliths reach up to 25 cm in diameter, show well-developed protuberances, and some of them are hollow (Fig. 7.7b). At 37 m water depth, the rhodolith coverage suddenly disappears. The coralline algae appearance is reduced to thin CAC attached to pebbles and cobbles which lie on the fine sediment, and Polysiphonia-like red algae appear. The coverage with Polysiphonia-like red algae increases with further seafloor shallowing and the CAC are reduced to growing on the upper parts of boulders (>1 m3), which occur occasionally and surmount the Polysiphonia fields. In 27 m

= JAGO dive track

= Dredge track

= Light measurements

684 = Station number 79°55‘

672 671 20

675 79°54‘ 685 depth [m] 684

1000 m

170 N 15°40‘ 15°50‘ E

Fig 7.6. Multibeam map of the Mosselbukta site showing depth and morphology of the seafloor and the positions of the applied gear (see Table 6.1 for a list of the kind and position of each applied gear); note the iceberg scouring-marks perpendicular to the shoreline; starting point of a track is always the numbered symbol.

42 Results water depth, two iceberg scouring-marks of c. 50 m in length appear. Here, the Polysiphonia- like red algae are removed and the remaining CAC are bleached. The marks carved into the seafloor and bleached, exhumed CAC and rhodoliths of unknown age are visible. Dive track 684 (78-44 m water depth), perpendicular to the northern Mosselbukta shoreline, shows a presumably glaciogenic pebble and cobble pavement. In 78 m water depth, living barnacles are very frequent and CAC are absent. The size of the cobbles increases from 74 m water depth and boulders (>1 m3) are visible. First CAC appear in 70 m water depth but are very rare (<5% coverage). In 64 m water depth, CAC on larger cobbles and boulders become more frequent (c. 10% coverage) and the first protuberances are developed. In 62 m water depth, a patch of pebbles and cobbles is covered with a fine sediment layer and CAC are not detectable. Development of protuberances markedly increases at a depth of 55 m, and CAC cover c. 30% of the larger cobbles. In 46 m water depth, rhodoliths up to 20 cm in diameter cover 60-80% of the seafloor, and many of the specimens are hollow. A small field of Polysiphonia-like red algae occurs at 46 m water depth, and CAC and rhodoliths disappear in the areas where these filamentous red algae grow. a

Fig. 7.7. JAGO-photographs of the seafloor at Mosselbukta; (a) well-developed rhodoliths in 42 m water depth, note the ophiurid arms between the rhodolith interstices; (b) partly hollow rhodoliths in 42 m water depth, note the cracked hollow rhodolith (arrow); scale bars = 10 cm.

43 Results

The Nordkappbukta site (80°31’N, 19°52’E) mapped by multibeam echosounder covers c. 16 km2. The seafloor has a distinct morphology that features zones of relatively flat depressions and plateaus (c. 30 m water depth) alternating with steep ridges, ribs and slopes down to c. 250 m water depth, while large iceberg scouring-marks are not detectable (Fig. 7.8). Dive track 701 (78-30 m water depth; see also Fig. 8.1a-c in the discussion chapter) perpendicular to the coast-loine in the eastern part of Nordkappbukta shows a bumpy, heavily struc- tured seafloor and heads along a slope. The inclination of the slope is c. 32° on average. The seafloor consists of a gravel pavement with poorly sorted material that is partly covered by a thin fine sediment layer, and some of the larger cobbles (>10 cm diameter) are covered with initial CAC. The CAC appear only if the substratum is clean, and are absent if it is smothered with fine sediment. At 75 m water depth, several large boulders (>1 m3) occur and are mostly completely covered with CAC (Fig. 7.9a). The boulders disappear at 70 m water depth and the seafloor shows the same appearance as in 78 m water depth, featuring patches with shell accumulations mainly from C. islandica and H. arctica. In 60 m water depth, the shell beds are absent and the seafloor is dominated by boulders and gravel, while the amount of CAC and their

= JAGO dive track

= Dredge track 80°33‘ = Light measurements

= CTD measurements

701 = Station number

702 706

701 80°32‘

711 712 703 20

704 depth [m] 80°31‘ 1000 m 250 N 19°40‘ 19°50‘ E Fig 7.8. Multibeam map of the Nordkappbukta site showing depth and morphology of the seafloor and the positions of the applied gear (see Table 6.1 for a list of the kind and position of each applied gear); starting point of a track is always the numbered symbol (after Teichert et al. 2012).

44 Results protuberance development increase. A steep slope at 51 m water depth made up by bedrock shows many CAC with well-developed protuberances and the first rhodoliths accumulated in cracks (Fig. 7.9b). The seafloor flattens at 45 m water depth and nearly the whole substratum made up by pebbles and cobbles is covered with CAC with well-developed protuberances and with rhodoliths up to 20 cm in diameter. The shell beds in this area consist mainly of C. islandica and H. arctica and are restricted to patches. A gravel pavement at 31 m water depth shows an increased coverage with Polysiphonia-like red algae whereas the rhodolith coverage decreases. Iceberg scouring-marks are absent along the entire track. Dive track 703 (95-110 m water depth) along a flat depression in the eastern part of Nordkapp bukta is very short and shows a pebble and cobble pavement with large boulders (>1 m3) on a smooth slope with c. 6° inclination. Epibenthos and shell accumulations are sparse and limited to depressions, where the pavement is also covered with a thin sediment layer. CAC and rhodoliths are absent. The seafloor at dive track 711 (70-30 m water depth; see also Fig. 8.1d-f in the discussion chapter) in the western part of Nordkappbukta mainly conforms to the observations from dive track 701, but the morphology of the seafloor is less pronounced, the total inclination of the slope is c. 15°, and bedrock is absent. The dive track starts on a gravel pavement with patchy distributed shell accumulations consisting mainly of C. islandica and H. arctica. Thin CAC appear on individual larger cobbles at 70 m water depth, and the frequency and thickness of the CAC increase with decreasing water depth. Occasionally, large boulders (>1 m3) occur and their upper parts are completely covered with CAC. In 50 m water depth, shell beds and boulders are absent and a flat gravel pavement dominates, while the bulk of this substrate is covered with CAC. Rhodoliths successively displace the coralline algal-encrusted cobble community at 45 m water depth and partly cover the seafloor (Fig. 7.10a). In 36 m water depth, boulders encrusted by CAC occur and a small area of the seafloor is affected by an iceberg scouring-mark leaving bleached rhodoliths and CAC (Fig. 7.10b). Polysiphonia-like red algae appear and increasingly cover the seafloor in 32 m water depth, so that the rhodoliths and CAC become successively replaced.

Appearance and properties of the rhodolith and CAC beds slightly vary along the different dive tracks at the four sites, but they show the same overall pattern. The initial growth of CAC starts at c. 78 m water depth if suitable substrate is available. With further shallowing of the seafloor, the development of the CAC protuberances continously increases, i.e. the protuberances are bigger and grow denser. The CAC coverage amounts up to 30% below c. 45 m water depth and the protuberances of the CAC are well-developed. This coincides with the first appearance of rhodoliths. With further flattening, the coverage with CAC and rhodoliths strongly increases up to 100% (with varying percentages of CAC and rhodoliths) if adequate substratum is available and if there is no shading and replacing by Polysiphonia-like red algae, whose appearance usually confines the coralline algae to a minimum water depth of c. 30 m.

45 Results a

Fig. 7.9. JAGO-photographs of the seafloor at Nordkappbukta; (a) boulders completely covered with CAC and living barnacles, sea squirts, and porifers in 75 m water depth; (b) bedrock covered with well-developed CAC, rhodoliths, living porifera and sea squirts in 45 m water depth; scale bars = 10 cm.

46 Results a

Fig. 7.10. JAGO-photographs of the seafloor at Nordkappbukta; (a) seafloor covered with well-developed CAC, rhodoliths, and shells of H. arctica (dead) in 45 m water depth; (b) iceberg-scouring mark with bleached CAC, rhodoliths, and shells of H. arctica (dead) in 36 m water depth; scale bars = 10 cm.

47 Results

7.2 Temperature and salinity patterns

The CTD records from stations 616 (Floskjeret), 632, 640, 641, 642, and 643 (Krossfjorden), 669 (Mosselbukta), and 699 and 706 (Nordkappbukta) show a similar pattern of increasing salinities and decreasing temperatures with depths (Fig. 7.11). This indicates a stratified summer situation which is characteristic for the coastal waters of Svalbard (Svendsen et al. 2002). The stations from Floskjeret and Krossfjorden, both being fjord localities, show a distinct -sur face layer with reduced salinity (c. 34) developed in the upper 10 m of the water column. This feature indicates an incumbent layer of brackish water that derives from the glaciers in the inner parts of the fjords. The discharged melt water has a lower density and mixes only successively with the underlying fully marine water masses. This is also the reason why the stratified pattern is very distinctive in the inner part of the fjords and becomes less pronounced to the fjord mouths. Underneath the incumbent layer of brackish water, the salinity increases continuously (>34.5), probably representing AW deriving from the WSC. At stations 669 (Mos- selbukta) and 699 and 706 (Nordkappbukta), this pattern is less developed since they are non-fjord localities and the mixing with more saline waters occurs much faster. However, the stratification is still visible on a smaller scale. The decrease of temperature with depth at the several CTD-measurement stations is not continuously but fluctuates such as it becomes particularly clear at station 616 in c. 120 m water depth (fluctuation between 2.0°C and 4.0°C) and at station 641 in c. 25 m water depth (fluctuation between 3.8°C and 5.3°C) . This characteristic arises due to the mixing of warm water masses deriving from the WSC with cold water masses from the ESC and holds for both, Isfjorden and Krossfjorden being fjord localities, and Mosselbukta and Nordkappbukta being non-fjord localities. This mixture develops turbulences that lead to in parts increasing temperatures with water depth (up to 2°C from c. 75 m to c. 100 m water depth in profile 642) where boundaries of different water masses are crossed by the CTD profile. Fig 7.12a; b shows salinity and temperature transects through Krossfjorden (see Fig. 7.3 for the transect course) computed by Ocean Data View 4 (Schlitzer 2012) using the data from the several CTD-profiles (stations 640, 641, 642, 643, and 632 from the inner fjord to the fjord mouth). The salinity transect shows the incumbent layer of brackish water deriving from the melt water discharge by the glacier, which becomes less pronounced towards the fjord mouth due to the successive mixing with the more saline underlying water layers. The temperature transect shows cold water masses from the ESC in the inner part of the fjord and warm water masses from the WSC at the fjord mouth. These water masses mix and develop turbulences which becomes particular clear at station 642, where the temperature pattern fluctuates with depth as described above (see also Fig. 7.11 for the CTD-profile). The observations, although being snapshots, clearly indicate the dynamic nature of the prevailing oceanographic regime. This regime strongly depends both on the annual changing characteristics of the ESC and the WSC, and the changing amounts of glacial melt water discharge as well.

48 Results 642 salinity (PSS) temperature [°C] temperature

0 1 2 3 4 5 6 7

31 32 33 34 35 36 20-0 10-0 5 0 -50 -100 -150 -200 -250 -300 -350 -400 depth [m] depth CTD-profiles from from 7.11. CTD-profiles Fig. 632, 616 (Floskjeret), stations - 643 (Kross 640, 641, 642, and 669 (Mosselbukta), fjorden), - 706 (Nordkapp 699 and and pat- similar a showing bukta) salinities increasing tern of - tempe decreasing and (blue) depths. with (red) ratures 641 706 salinity (PSS) salinity (PSS) temperature [°C] temperature temperature [°C] temperature

0 1 2 3 4 5 6 7 0 1 2 3 4 5 6 7

31 32 33 34 35 36 31 32 33 34 35 36 20-0 10-0 5 0 -50 -100 -150 -200 -250 -300 -350 -400 20-0 10-0 5 0 -50 -100 -150 -200 -250 -300 -350 -400

depth [m] depth depth [m] depth 699 640 salinity (PSS) salinity (PSS) temperature [°C] temperature temperature [°C] temperature

0 1 2 3 4 5 6 7 0 1 2 3 4 5 6 7

31 32 33 34 35 36 31 32 33 34 35 36 20-0 10-0 5 0 -50 -100 -150 -200 -250 -300 -350 -400 20-0 10-0 5 0 -50 -100 -150 -200 -250 -300 -350 -400

depth [m] depth depth [m] depth 632 669 salinity (PSS) salinity (PSS) temperature [°C] temperature temperature [°C] temperature

0 1 2 3 4 5 6 7 0 1 2 3 4 5 6 7

31 32 33 34 35 36 31 32 33 34 35 36 20-0 10-0 5 0 -50 -100 -150 -200 -250 -300 -350 -400 20-0 10-0 5 0 -50 -100 -150 -200 -250 -300 -350 -400

depth [m] depth depth [m] depth 616 643 salinity (PSS) salinity (PSS) temperature [°C] temperature [°C] temperature

0 1 2 3 4 5 6 7 0 1 2 3 4 5 6 7

31 32 33 34 35 36 31 32 33 34 35 36 20-0 10-0 5 0 -50 -100 -150 -200 -250 -300 -350 -400 20-0 10-0 5 0 -50 -100 -150 -200 -250 -300 -350 -400

depth [m] depth [m] depth

49 Results

a CTD - station 632 643 642 641 640 35

35 depth [m] 200 100 0 salinity (PSS)

36 300

37 0 10 20 30 40 50 distance [km]

b CTD - station 632 643 642 641 640 7

3 depth [m] 200 100 0 temperature [°C] temperature 2

1 300

0 0 10 20 30 40 50 distance [km]

Fig. 7.12. CTD-transects through Krossfjorden from the fjord mouth (left) to the glacier slope of Lilliehöökbreen (right; see Fig. 7.3 for the transect course); (a) decreasing salinity towards the inner part of the fjord due to the successive mixing of saline water masses with an incumbent layer of brackish water deriving from the glacier slope; (b) increasing mixture of the cold ESC fjord water with the warmer AW from the WSC towards the fjord mouth; transects were created with Ocean Data View 4 (Schlitzer 2012).

50 Results

7.3 The light regime

The photosynthetically active radiation (PAR; unit = µmol photons -2m s-1) was measured under various weather conditions in order to determine the depths of the photic zoning. The surface illumination at stations 654 (Krossfjorden), 672/1 and 672/2 (Mosselbukta), and 702 and 712 (Nordkappbukta) ranged from 636 to 2580 µmol photons m-2 s-1 above and from 218 to 1052 µmol photons m-2 s-1 just below the water surface, and the light levels decreased exponentially with water depth. No light measurements were done at Floskjeret. To compare the absolute PAR-levels of the study sites, the data were plotted on an absolute and on a logarithmic scale (Fig. 7.13). For the determination of the boundaries between the photic zones, the data were plotted on a logarithmic scale with respect to the previously measured surface illumination. The lower boundary of the euphotic zone (1% surface illumination) is at c. 45 m water depth at stations 654, 702 and 712, and at 20-25 m water depth at stations 672/1 and 672/2. The lower boundary of the dysphotic zone (0.01% surface illumination) is only reached in profile 672/2, in a water depth of c. 64 m. At the Krossfjorden site (station 654), the decrease of the PAR processes remarkably faster than at the Mosselbukta and Nordkappbukta sites, due to the considerable amount of melt water discharge by the glacier at the end of the fjord. This observation is in accordance to the observations made while descending during the JAGO dives.

51 Results

a PAR [µmol photons m-2s-1] station 654 PAR [µmol photons m-2s-1] station 654 % surface illumination station 654 0 200 400 600 800 1000 1200 0 0.01 0.1 1 10 100 1000 10000 0 0.01 0.1 1 10 100 0 0 0 10 10 10 20 20 20 30 30 30 40 40 40 50 50 50 60 60 60 depth [m] depth [m] depth [m] 70 70 70 80 80 80 90 90 90 aphotic dysphotic euphotic 100 100 100

station 672/1 station 672/1 station 672/1 b PAR [µmol photons m-2s-1] PAR [µmol photons m-2s-1] % surface illumination station 672/2 station 672/2 station 672/2 0 200 400 600 800 1000 1200 0 0.01 0.1 1 10 100 1000 10000 0 0.01 0.1 1 10 100 0 0 0 10 10 10 20 20 20 30 30 30 40 40 40 50 50 50 60 60 60 depth [m] depth [m] depth [m] 70 70 70 80 80 80 90 90 90 aphotic dysphotic euphotic 100 100 100

station 702 station 702 station 702 c PAR [µmol photons m-2s-1] PAR [µmol photons m-2s-1] % surface illumination station 712 station 712 station 712 0 200 400 600 800 1000 1200 0 0.01 0.1 1 10 100 1000 10000 0 0.01 0.1 1 10 100 0 0 0 10 10 10 20 20 20 30 30 30 40 40 40 50 50 50 60 60 60 depth [m] depth [m] depth [m] 70 70 70 80 80 80 90 90 90 aphotic dysphotic euphotic 100 100 100 Fig. 7.13. Measurements of the light regime at (a) Krossfjorden (station 654); (b) Mosselbukta (stations 671/1 and 672/2); (c) Nordkappbukta (stations 702 and 712); the diagrams from right to left show PAR on an absolute scale, PAR on a logarithmic scale, and PAR with respect to surface illumination indicating the photic zoning.

52 Results

7.4 Chemistry of the seawater

The water samples to measure total alcalinity (TA), dissolved inorganic carbon (DIC), temperature, and salinity were collected by the bottles mounted to the CTD rosette and by the Niskin bottle attached to the JAGO submersible.

The carbonate system parameters pH, carbon dioxide partial pressure (pCO2), calcite saturation (ΩCal) and aragonite saturation (ΩArg) were calculated from TA, DIC, temperature and salinity. All values are compiled in Table 7.1. The temperatures are relatively low indicating that most stations are under the influence of the ESC. Due to the great water depth, the salinities show no influence of an incumbent layer of brackish water but indicate fully marine conditions.

The pH is >8 and pCO2 is <390 µatm for all water depths where rhodoliths occur. Despite the low temperatures, ΩCal is >1 and thus in the range of saturation at all stations. The same applies for ΩArg except for stations 616 and 632, which are slightly undersaturated with respect to aragonite but were measured in water depths where rhodoliths do not occur.

Table 7.1. Water chemistry measurements of the four sites; T = temperature; p = pressure; TA = total alkalinity; DIC = dissolved inorganic carbon; pCO2 = CO2 partial pressure ΩCal = calcite saturation;

ΩArg = aragonite saturation.

station # T p salinity TA DIC pH pCO2 ΩCal ΩArg [°C] [dbars] (PSS) [µmol/kgSW] [µmol/kgSW] (total scale) [µatm] 616 0.8 417 34.8 2292 2227 7.85 593 1.44 0.91 632 1.3 332 34.8 2244 2178 7.85 589 1.45 0.92 640 1.4 132 34.5 2288 2125 8.14 298 2.76 1.74 641 1.1 231 34.7 2274 2144 8.06 363 2.26 1.42 642 0.2 341 34.9 2312 2215 7.96 458 1.81 1.14 643 1.3 283 34.8 2288 2148 8.07 348 2.36 1.49 652 3.0 47 35.0 2252 2098 8.09 336 2.67 1.68 684 3.0 44 35.0 2283 2136 8.07 360 2.59 1.64 699 3.0 131 35.0 2285 2106 8.15 294 2.97 1.88 703 3.0 30 35.0 2262 2096 8.12 315 2.84 1.79 706 3.7 62 34.6 2276 2110 8.11 324 2.86 1.80 714 3.0 50 35.0 2266 2084 8.16 285 3.06 1.93

53 Results

7.5 Composition of the benthic community

A total of 41 (Krossfjorden), 42 (Mosselbukta), and 42 (Nordkappbukta) benthic species (or higher taxa) were identified in the dredge samples and from the JAGO video footage. No dredge samples were taken at the Floskjeret site. These numbers, especially those gained during the JAGO dives, are considered as conservative measures of species diversity regarding the higher amount of taxa that can be found when a larger variety of sampling tools is used (Hall-Spen- cer & Atkinson 1999). Due to the influence of the North Atlantic Current system featuring the warm AW which is transported northwards by the WSC, most benthic species inhabiting the coastal waters of the Svalbard archipelago, especially at the western and northern coasts, are actually of Atlantic origin, i.e. they do also occur at boreal latitudes. Endemic Arctic species adapted to extremely low water temperatures, as they characterise polar environments and do occur in Arctic waters only, were not identified . The percentaged fractions regarding the higher taxonomic levels are shown in Fig. 7.14, and the taxa identified at each site (excluding Floskjeret in each case) are compiled in Table 7.2. The important grazers Tonicella rubra and Strongylocentrotus sp. are present on rhodoliths from all sites, including Floskjeret (see Fig. 7.15 and 7.16 for the most prominent animal species inhabiting the rhodolith beds). Where rhodoliths are present, many benthic organisms can be found on the surfaces of the rhodoliths, in interstices between them and inside hollow rhodoliths (see Table 7.3 for the different ways of interrelation between the benthos and the rhodolith beds). In this coherence, the CT-scans of the rhodoliths from the Mosselbukta site from 44 m water depth clearly show that the rhodoliths contain many associated organisms (Fig. 7.17). Some organisms like the serpulids (Fig. 7.17c; f-h) occur almost exclusively inside of hollow rhodoliths. Krossfjorden exhibits the lowest species numbers of the three investigated sites with respect to the benthic community. Regarding species numbers, Crustacea dominate with 13 identified taxa (31%) and are followed by Echinodermata with 11 identified taxa (26%) and Pisces with 6 identified taxa (14%). Mosselbukta features a more balanced species assemblage, with Mol- lusca and Echinodermata exhibiting the highest numbers of identified taxa (8 respectively 19% each). At Nordkappbukta, the species assemblage is also balanced with most identified taxa for Mollusca (9 respectively 22%) and Echinodermata (8 respectively 19%). The ways of interrelation between the identified taxa and the rhodolith beds are manifold (Table 7.3) and specified as follows. The sponge genus Geodia sp. was identified from Nordkappbukta and overgrows the coralline algal crusts, being a competitor for suitable substratum. Various fixosessile cnidarians (Actinia sp., Gersemia rubiformis, and Hormathia nodosa at Nord- kappbukta and Lucernaria quadricornis at Krossfjorden and Mosselbukta) are fixed to the surface of rhodoliths and act as competitors for space. Many molluscs are in close interrelationship with the rhodoliths, especially the Iceland scallop (Chlamys islandica; Fig. 7.15d) and the wrinkled rockborer (Hiatella arctica; Fig. 7.15e), which can be found inside hollow rhodoliths and, in the case of H. arctica, also in the calcitic matrix of the coralline algae (see also Fig. 7.17a-e). Chlamys islandica and H. arctica were identified at all sites. The crenulate astarte (Astarte crenata; Fig. 7.15 c) could also

54 Results be found in hollow rhodoliths and was identified at Nordkappbukta. Of great importance for the rhodoliths are the grazing molluscs, from which the northern red chiton (Tonicella rubra; Fig. 7.15g) is the most abundant and could be found at all sites, where it was feeding on the rhodolith surfaces. This feeding behaviour also holds for Noah’s keyhole limpet (Punctu- rella noachina), that was identified at Mosselbukta and Nordkappbukta, and for Tectura sp. (Fig. 7.15f), that was identified at Nordkappbukta. Altogether, nine different taxa of polychaetes were identified, with Thelepus cincinnatus being the only species identified at Krossfjorden, Mosselbukta, and Nordkappbukta. The polychaetes occur to live inside hollow rhodoliths and also in the interstices between them. The calcareous tubes of non further determined polychaetes were found to be almost completely restricted to the inside of hollow rhodoliths (see also Fig. 7.17f-h). Pantopoda were identified from Mosselbukta and Nordkappbukta but showed no way of interrelation with the rhodolith beds. Crustaceans are abundant at all sites, but the only interrelations to the rhodoliths were observed for the hermit crab Pagurus pubescens, that is frequently present in the interstices between individual rhodoliths, and specimens of Balanus sp. (Fig. 7.15i; Fig. 7.17g-h), that grow on the rhodoliths surfaces and act as competitors for space. On the other hand, many nektonic crustaceans (e.g. Lebbeus polaris, identified from all sites; Fig. 7.16a) were visible just above the rhodolith beds, so an interrelation cannot be completely excluded. Bryozoans could be found at all sites. Cellepora sp. and Myriapora coarctata were identified at Krossfjorden and Mosselbukta and Flustra foliaceae (Fig. 7.15a) at Nordkappbukta. They grow on the rhodoliths surfaces and hence act as competitors for space. The only identified brachiopod species is Hemithiris psittacea (Fig. 7.15b), that rarely occurred at Mosselbukta and Nordkappbukta, where it could been found attached to the surface of rhodoliths. The Echinodermata are the group with most identified taxa (14) in relation to the total sum of all sites. The echinoderms most closely related to the rhodoliths are the sea urchins Strongylo- centrotus sp. (identified from all sites; Fig. 7.16g) and Strongylocentrotus pallidus (identified from Krossfjorden), since they are heavily feeding on the rhodoliths surfaces and act as bioeroders. Brittlestars are very common, especially Ophiacantha bidentata (Fig. 7.16c) and Ophi- ura robusta (Fig. 7.16d), which were identified from all sites. The brittlestars use the interstices between the rhodoliths as shelter and are collecting detritus with their arms (see also Fig. 7.7a). Starfish like Henricia sanguinolenta (identified at Mosselbukta and Nordkappbukta; Fig. 7.16b) also use the interstices between the rhodoliths as shelter. The fixosessile sea squirt species Boltenia echinata (Fig. 7.16e) and Styela rustica (Fig. 7.16f) were identified from Mosselbukta and Nordkappbukta. They settle also on the surface of rhodoliths but were not very common. Fish were numerous, but except for the Atlantic hookear sculpin (Artediellus atlanticus) and the snake blenny (Lumpenus lampretaeformis), which use the interstices between the rhodoliths as shelter, no interrelation with the rhodoliths could be observed. In this coherence one has to mention the first sighting of the Atlantic snake pipefish (Entelurus aequoreus) at Krossfjorden, a species that was presumed to be confined to the area south of Iceland (Fleischer et al. 2007).

55 Results

Rhodophyta Cnidaria 2% Rhodophyta 2% 5% Cnidaria Ascidiacea 3% 7% Pisces Pisces 7% 14% Mollusca 10%

Mollusca Polychaeta 19% 10% Echinodermata Echinodermata 19% 26% Polychaeta 14% Crustacea 31% Brachiopoda 2% Bryozoa Crustacea 5% 14% Bryozoa Pantopoda 5% 5% Kross orden Mosselbukta

Pisces 2%

Ascidiacea Rhodophyta 5% 7% Porifera 3%

Cnidaria 7% Echinodermata 19%

Brachiopoda 2% Mollusca 22%

Bryozoa 2% Crustacea 17% Polychaeta 12%

Pantopoda 2% Nordkappbukta

Fig. 7.14. Pie charts showing the pecentages of higher taxonomic levels at Krossfjorden, Mosselbukta and Nordkappbukta. Krossfjorden has a lower number of species than Mosselbukta and Nordkapp- bukta, and is dominated by Crustacea, Echinodermata and Pisces.

56 Results

Table 7.2. Presence and absence data of the identified taxa at each site (excluding Floskjeret).

Taxon Taxon Krossfjorden Mosselbukta Nordkappbukta Krossfjorden Mosselbukta Nordkappbukta

Rhodophyta Crustacea ff.

Lithothamnion glaciale Kjellman, 1883 XXX Spirontocaris phippsii (Krøyer, 1841) X

Phymatolithon tenue (Rosenvinge) Düwel & Wegeberg, 1996 X Spirontocaris spinus (Sowerby, 1805) XXX

Polysiphonia-like red algae XX Bryozoa

Porifera Cellepora sp. Linnaeus, 1767 XX

Geodia sp. Lamarck, 1815 X Flustra foliacea (Linnaeus, 1758) X

Cnidaria Myriapora coarctata (Sars, 1863) XX

Actinia sp. Linnaeus, 1776 X Brachiopoda

Gersemia rubiformis (Ehrenberg, 1834) X Hemithiris psittacea (Gmelin, 1790) XX

Hormathia nodosa (Fabricius, 1780) X Echinodermata

Lucernaria quadricornis Müller, 1776 XX Crossaster papposus (Linnaeus, 1767) XX

Mollusca Gorgonocephalus sp. Leach, 1815 X X

Astarte crenata (Gray, 1824) X Heliometra glacialis (Owen, 1833) X X

Boreotrophon clathratus (Linnaeus, 1767) X Henricia sanguinolenta (Müller, 1776) XX

Boreotrophon truncatus (Ström, 1768) XX Leptasterias sp. Verrill, 1866 X

Buccinum sp. Linnaeus, 1758 X Ophiacantha bidentata (Bruzelius, 1805) XXX

Cadlina laevis (Linnaeus, 1767) X Ophiocten sericeum (Forbes, 1852) X X

Chlamys islandica (Müller, 1776) XXX Ophiopholis aculeata (Linnaeus, 1767) XXX

Hiatella arctica (Linnaeus, 1767) XXX Ophiura robusta (Ayres, 1854) XXX

Musculus laevigatus (Gray, 1824) X Ophiura sarsii Lütken, 1855 X

Puncturella noachina (Linnaeus, 1771) XX Pteraster militaris (Müller, 1776) X

Tectura sp. Gray, 1847 X Solaster endeca (Linnaeus, 1771) X

Tonicella rubra (Linnaeus, 1767) XXX Strongylocentrotus sp. Brandt, 1835 XXX

Tridonta montagui (Dillwyn, 1817) X Strongylocentrotus pallidus (Sars, 1871) X

Velutina sp. Fleming, 1820 X Ascidiacea

Polychaeta Boltenia echinata (Linnaeus, 1767) XX

Brada sp. Stimpson, 1854 X Styela sp. Fleming, 1822 X

Eunoe nodosa (Sars, 1861) X Styela rustica Linnaeus, 1767 XX

Flabelligera affinis Sars, 1829 XX Pisces

Harmothoe imbricata (Linnaeus, 1767) X Artediellus atlanticus Jordan & Evermann, 1898 XX

Nephtys sp. Cuvier, 1817 X X Entelurus aequoreus (Linnaeus, 1758) X

Nereis zonata Malmgren, 1867 XX Eumicrotremus spinosus (Fabricius, 1776) X

Nothria conchylega (Sars, 1835) X Gadus morhua Linnaeus, 1758 X

Terebellides stroemii Sars, 1835 XX Liparis sp. Scopoli, 1777 XX

Thelepus cincinnatus (Fabricius, 1780) XXX Lumpenus lampretaeformis (Walbaum, 1792) X

Pantopoda Melanogrammus aeglefinus (Linnaeus, 1758) X

Nymphon sp. Fabricius, 1794 XX

Phoxichilidium femoratum (Rathke, 1799) X

Crustacea

Anonyx laticoxae Gurjanova, 1962 X X

Balanus sp. Costa, 1778 XXX

Hyas araneus (Linnaeus, 1758) XXX

Hyas coarctatus Leach, 1816 X

Lebbeus polaris (Sabine, 1824) XXX

Monoculodes borealis Boeck, 1871 X

Mysis oculata (Fabricius, 1780) X

Pagurus pubescens Krøyer, 1838 XXX

Sabinea sarsii Smith, 1879 X

Sabinea septemcarinata (Sabine, 1824) X

Sclerocrangon ferox (Sars, 1877) XXX

57 Results

a b c

Fig. 7.15. Prominent epibenthos inhabiting the rhodolith beds; (a) Flustra foliacea with serpulids; (b) Hemithiris psittacea; (c) Astarte crenata; (d) Chlamys islandica; (e) Hiatella arctica; (f) Tectura sp.; (g) Tonicella rubra; (h) Polychaete, suborder Terebellomorpha; (i) Balanus sp.; scale bars = 0.5 cm (slightly modified after Teichert et al. 2012).

58 Results

g f

Fig. 7.16. Prominent epibenthos inhabiting the rhodolith beds; (a) Lebbeus polaris; (b) Henricia sanguinolenta; (c) Ophiacantha bidentata; (d) Ophiura robusta; (e) Boltenia echinata; (f) Styela rustica; (g) Strongylocentrotus sp.; scale bars = 0.5 cm (slightly modified after Teichert et al. 2012).

59 Results

Table 7.3. List of the identified taxa and their interrelations to the rhodolith beds.

Interrelation to rhodolith beds Interrelation to rhodolith beds

Species Species no observations living on rhodoliths surface rhodoliths surface to attached rhodoliths living inside hollow rhodoliths inside hollow attached on rhodoliths surfacefeeding rhodolithsliving between rhodoliths overgrowing no observations living on rhodoliths surface rhodoliths surface to attached rhodoliths living inside hollow rhodoliths inside hollow attached on rhodoliths surfacefeeding rhodolithsliving between rhodoliths overgrowing

Rhodophyta Crustacea ff.

Lithothamnion glaciale X Monoculodes borealis X

Phymatolithon tenue X Mysis oculata X

Polysiphonia-like red algae X Pagurus pubescens X

Porifera Sabinea sarsii X

Geodia sp. X X Sabinea septemcarinata X

Cnidaria Sclerocrangon ferox X

Actinia sp. X Spirontocaris phipsii X

Gersemia rubiformis X Spirontocaris spinus X

Hormathia nodosa X Bryozoa

Lucernaria quadricornis X Cellepora sp. X

Mollusca Flustra foliacea X X

Astarte crenata X X Myriapora coarctata X

Boreotrophon clathratus X Brachiopoda

Boreotrophon truncatus X Hemithiris psittacea X

Buccinum sp. X Echinodermata

Cadlina laevis X Crossaster papposus X

Chlamys islandica X X Gorgonocephalus sp. X

Hiatella arctica X X Heliometra glacialis X

Musculus laevigatus X Henricia sanguinolenta X

Puncturella noachina X X Leptasterias sp. X

Tectura sp. X X Ophiacantha bidentata X

Tonicella rubra X X Ophiocten sericeum X X

Tridonta montagui X Ophiopholis aculeata X

Velutina sp. X Ophiura robusta X X

Polychaeta Ophiura sarsii X X

Brada sp. X Pteraster militaris X

Eunoe nodosa X Solaster endeca X

Flabelligera affinis X Strongylocentrotus sp. XX

Harmothoe imbricata X Strongylocentrotus pallidus XX

Nephtys sp. X Ascidiacea

Nereis zonata X X Boltenia echinata X

Nothria conchylega X Styela sp. X

Terebellides stroemii X X Styela rustica X

Thelepus cincinnatus X Pisces

Pantopoda Artediellus atlanticus X

Nymphon sp. X Entelurus aequoreus X

Phoxichilidium femoratum X Eumicrotremus spinosus X

Crustacea Gadus morhua X

Anonyx laticoxae X Liparis sp. X

Balanus sp. X Lumpenus lampretaeformis X

Hyas araneus X Melanogrammus aeglefinus X

Hyas coarctatus X

Lebbeus polaris X

60 Results

a b

C. islandica

H. arctica

c d

H. arctica C. islandica

serpulids

lithoclast e f

serpulids ophiurid

H. arctica

g barnacle h

barnacles serpulids

serpulids

ophiurid

Fig. 7.17. CT-scans of rhodoliths from Mosselbukta at 44 m water depth showing various occupying organisms; (a) C. islandica, note the conductor muscle in lower right shell; (b) H. arctica; (c) C. islandica and serpulids; (d) H. arctica overgrown by L. glaciale, note embodied lithoclast; (e) H. arctica and ophiurid; (f) serpulids; (g) barnacle, serpulids, and ophiurid; (h) serpulids and barnacles; scale bars = 1 cm.

61 Results

7.6 Taxonomy of the coralline algae

The coralline algal rhodolith community of Floskjeret is composed of Lithothamnion glaciale Kjellman, 1883 (Fig. 7.18a-d) and Phymatolithon tenue (Rosenvinge) Düwel & Wege- berg, 1996 (Fig. 7.19a-c) in equal parts. The coralline algal rhodolith communities of Kross- fjorden and Mosselbukta are composed of L. glaciale only. The coralline algal rhodolith community of Nordkappbukta is mainly composed of L. glaciale, with a lesser amount of P. tenue.

Lithothamnion glaciale is classified as follows:

Empire Eukaryota Chatton, 1925 Kingdom Plantae Haeckel, 1866 Subkingdom Biliphyta Cavalier-Smith, 1981 Phylum Rhodophyta Wettstein, 1901 Subphylum Eurhodophytina Saunders & Hommersand, 2004 Class Florideophyceae Cronquist, 1960 Subclass Corallinophycidae Cronquist, 1960 Order Corallinales Silva & Johansen, 1986 Family Hapalidiaceae Gray, 1864 Subfamily Melobesioideae Bizzozero, 1885 Genus Lithothamnion Heydrich, 1897 Species Lithothamnion glaciale Kjellman, 1883

The identification of thalli of Lithothamnion glaciale was based on data in Adey (1964; 1966; 1970b, including species keys), Adey et al. (2005, including comparisons of L. glaciale with Lithothamnion tophiforme (Esper) Unger, 1858 and Lithothamnion lemoineae Adey, 1970), Irvine & Chamberlain (1994, including species keys and accounts), and Kjellman (1883; 1885, containing the original account of L. glaciale). The living thalli of L. glaciale (Fig. 7.18a; b) have a dull texture and reddish to dull pink colour. The conceptacles that bear the tetra- spores have up to 50 pores in their roofs (Fig. 7.18c). Like other species of Lithothamnion, the vegetative filaments of L. glaciale are terminated by epithallial cells with flared outer corners (Fig. 7.18d). These characters distinguish L. glaciale from another Arctic species, Lithotham- nion tophiforme, that was not identified in the samples. Living thalli of L. tophiforme are characterised by a glossy texture, an orange-red colour, and tetrasporangial conceptacles with up to 85 pores in their roofs (Adey et al. 2005). Lithothamnion tophiforme is deemed to be primarily an Arctic species that extends only into the deeper and colder parts of the subarctic Atlantic (Adey et al. 2005), while L. glaciale is known as the dominant subarctic species of the genus Lithothamnion. Adey & Adey (1973) state that L. glaciale is the primary subarctic

62 Results rhodolith builder, partly being replaced by L. tophiforme in Arctic waters. However, the type specimen of L. glaciale comes from Spitsbergen, Svalbard, and in the original account of the species, Kjellman (1883) states that L. glaciale is common and plentiful on the west and north coasts of Spitsbergen and occurs as far north as Treurenberg Bay (79°56’N). According to Hansen & Jenneborg (1996: 372), L. glaciale is the dominant non-geniculate coralline algae at Spitsbergen. At Floskjeret, Krossfjorden, and Mosselbukta, L. glaciale grows as unattached rhodoliths and also attached to cobbles. At Nordkappbukta, L. glaciale additionally grows attached to bedrock. The rhodoliths are more or less spherical to ovoidal to more ir- regular in form, consist largely of knobby protuberances (branches), and are nucleated or hollow. Attached thalli produce short (up to 19 mm) warty or knobby protuberances of varying diameter from a crustose base.

a b

Fig. 7.18. Lithothamnion glaciale Kjellman, 1883; (a) group of living rhodoliths, most of which are hollow; scale bar = 5 cm; (b) individual rhodolith; scale bar = 1 cm; (c) tetrasporangial conceptacle in surface view, note conceptacle pores on flattened part of roof; scale bar = 100 µm; (d) section of thallus showing calcified filaments terminating in flattened and flared epithallial cells, note fusions between some cells of adjacent filaments; scale bar = 10 µm (modified after Teichert et al. 2012).

63 Results

Phymatolithon tenue is classified as follows:

Subfamily Melobesioideae Bizzozero, 1885 Genus Phymatolithon Foslie, 1898 Species Phymatolithon tenue (Rosenvinge) Düwel & Wegeberg, 1996

The identification of specimens of Phymatolithon tenue was based on Düwel & Wegeberg (1996) and Rosenvinge (1893). Phymatolithon tenue was originally described under the name Lithothamnion tenue by Rosenvinge (1893), but a modern examination of the lecto- type specimen by Düwel & Wegeberg (1996) showed that the species belongs to the genus Phymatolithon. The type specimen of P. tenue comes from Holsteinborg, Greenland. Phyma- tolithon tenue characteristically has a thin encrusting thallus that usually grows up to 200 µm in thickness (Fig. 7.19a; b). The thallus does not produce protuberances and has sporangial conceptacles that appear as white dots on the upper thallus surface (Fig. 7.19a). The pore

a b

Fig. 7.19. Phymatolithon tenue (Rosenvinge) Düwel & Wegeberg, 1996; (a) part of a living thallus, the white dots on the upper surface denote the positions of individual sporangial conceptacles; scale bar = 5 mm; (b) longitudinal section of the thallus edge showing a ventral core of filaments portions which curve upwards to the dorsal thallus surface; scale bar = 100 µm; (c) part of a tetrasporangial conceptacle roof showing three pore canals still blocked by sporangial plugs, note that cells lining the pore canals characteristically are somewhat swollen compared with other roof cells; scale bar = 25 µm (modified after Teichert et al. 2012).

64 Results canals of the conceptacles are lined with cells that are mostly slightly larger than other cells in the conceptacle roof (Fig. 7.19c). These features were described and illustrated by Düwel & Wegeberg (1996: 478, Figs. 29-33) in the type specimen and are clearly evident in specimens from Floskjeret and Nordkappbukta, where thalli of P. tenue grow attached to cobbles or, in the case of Nordkappbukta, even to bedrock, and occur intermixed with L. glaciale. During this study, many SEM pictures were taken in order to determinate the taxonomy of the coralline algae prevailing at the different sites. Some of these pictures also show the starch grains mentioned above in thalli of L. glaciale (Fig. 7.20) and hence indicate that the algae are well adapted to the demands of the polar night. The starch grains were not detected in SEM pictures of thalli of P. tenue, but testing the thalli with iodine indicates the presence of starch reservoirs due to the blue staining of the thalli (pers. obs.). The SEM pictures also revealed dense biofilms mainly consisiting of diatoms which are present on the rhodoliths (Fig. 7.21).

Fig. 7.20. Coloured SEM image showing a part of the perithallial tissue of L. glaciale from the Nord- kappbukta site. The starch grains are clearly visible inside the cell cavities and act as carbon reservoirs; scale bar = 5 µm.

65 Results

Fig. 7.21. SEM image showing a biofilm mainly consisting of diatoms on the epithallial tissue of L. glaciale from Nordkappbukta in 38 m water depth; scale bar = 5 µm.

66 Results

7.7 Description and classification of the rhodoliths

The following analyses to gain information about the shape and size ranges of the rhodoliths are based on specimens collected from 45 m water depth at Floskjeret, 41-50 m water depth at Krossfjorden, 44 m water depth at Mosselbukta, and 27-45 m water depth at Nordkappbukta. The rhodolith shape was analysed by measuring the long (L), intermediate (I) and short (S) axes of the specimens and applying the maximum projection sphericity formula [(S2/LI)0.5] as described in Bosence (1983a), and the results were plotted into a pebble shape diagram (Sneed & Folk 1958) using the MICROSOFT EXCEL® sheet Tri-plot (Graham & Midgley 2000) (Fig. 7.22). The rhodoliths from Floskjeret (n = 37) show a slight concentration on discoidal (43%) and spheroidal (38%) shapes, while ellipsoidal (19%) shapes are less abundant. The Krossfjorden rhodoliths

a Spheroidal b Spheroidal 1.0 1.0

0.9 0.9

0.8 0.8

0.7 0.7

0.6 0.6

(S/L) 0.5 (S/L) 0.5

0.4 0.4

0.3 0.3

0.2 0.2

0.1 0.1 Floskjeret Kross orden 0.0 0.0 0.0 0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8 0.9 1.0 0.0 0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8 0.9 1.0 Discoidal (L-I)/(L-S) Ellipsoidal Discoidal (L-I)/(L-S) Ellipsoidal

c Spheroidal d Spheroidal 1.0 1.0

0.9 0.9

0.8 0.8

0.7 0.7

0.6 0.6

(S/L) 0.5 (S/L) 0.5

0.4 0.4

0.3 0.3

0.2 0.2

0.1 0.1 Mosselbukta Nordkappbukta 0.0 0.0 0.0 0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8 0.9 1.0 0.0 0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8 0.9 1.0 Discoidal (L-I)/(L-S) Ellipsoidal Discoidal (L-I)/(L-S) Ellipsoidal

Fig. 7.22. Tri-plot shape distribution (Graham & Midgley 2000) of rhodoliths from the particular sites based on measurements of the long (L), intermediate (I) and short (S) axes as described in Bo- sence (1983a); (a) Floskjeret, n = 37; (b) Krossfjorden, n= 122; (c) Mosselbukta, n = 92; (d) Nord- kappbukta, n = 128.

67 Results

(n = 122) are dominated by spheroidal (58%) and ellipsoidal (29%) shapes, the rhodoliths from Mosselbukta (n = 92) were mostly of spheroidal shape (79%), and the rhodoliths from Nordkap- pbukta (n = 128) show a majority of spheroidal shapes (60 %), while the remaining rhodoliths disperse on discoidal and ellipsoidal shapes in c. equal parts (17% and 13%). The size of the rhodoliths was measured using the volume of an ellipsoid [(LIS/4π)0.5] as shown in Bosence (1976). Plotting the values into a box plot (Fig. 7.23) indicates that the size ranges of rhodoliths from Floskjeret (c. 2-11 cm), Mosselbukta (c. 1-12 cm), and Nordkappbukta (c. 1-13 cm) are much larger than those from Krossfjorden (c. 1-4 cm). The rhodolith community from Floskjeret is composed of 86% monospecific and 14% multispecific rhodoliths. Monospecific rhodoliths consist either of L. glaciale or P. tenue, while both species provide c. 50% of the overall surface coverage at Floskjeret each. Rhodoliths from Kross- fjorden and Mosselbukta are monospecific and consist of L. glaciale only. Most rhodoliths from Nordkappbukta are monospecific (85%) and consist of L. glaciale. Some are multispecific (15%) and both L. glaciale and P. tenue overgrow each other. Overall, L. glaciale provides about 90% of the surface coverage at Nordkappbukta. Both coralline algal species, L. glaciale and P. tenue, settle on hard substratum above a distinctive size that provides enough stability (Fig. 7.24). If bedrock is present as it holds for the Nordkapp- bukta site, it is also covered mainly by coralline algae, prior to porifers, barnacles, and sea squirts (Fig. 7.25). The rhodoliths generally contain a lithoclastic core and occur both mixed with the encrusting coralline algal cobble communities and as distinct accumulations. Hollow rhodoliths with a wide-open space for internal colonisation and settlement to form a specific cryptic mi- crohabitat also occur (Fig. 7.26, see also CT-scans in Fig. 7.17), except for the Krossfjorden site. Although the system of their development is not yet fully understood, it seems that the hollow rhodoliths have lost their lithoclastic nucleus at an earlier growth stage.

size [cm] size 6

0 a b c d

Fig. 7.23. Box plots showing the size ranges of Fig. 7.24. Coralline algal crusts of different sta- rhodoliths from the particular sites using the vo- ges of development covering cobbles on a gravel lume of an ellipsoid [(LIS/4π)0.5] as described in flat at 70 m water depth at Nordkappbukta; scale Bosence (1976); (a) Floskjeret, n = 37; (b) Kross- bar = 5 cm. fjorden, n = 122; (c) Mosselbukta, n = 92; (d) Nord- kappbukta, n = 128.

68 Results

Fig. 7.25. Well-developed coralline algal crusts Fig. 7.26. Hollow rhodolith containing a Chla- nearly entirely covering the bedrock at 45 m wa- mys islandica specimen from 42 m water depth ter depth at Nordkappbukta; scale bar = 5 cm at Mosselbukta; scale bar = 1 cm.

69 Results

7.8 Verification of the annual rhodolith banding pattern

Longitudinal sections through the protuberances of Lithothamnion glaciale show a distinct pattern of growth increments, at which each increment consists of a varying number of calcified cell rows. The increments start with dense, heavily calcified cell rows (i.e. there are no or only small cell cavities left), grading into less calcified cell rows (i.e. there are large cell cavities left), and each increment sequence is presumed to represent one annual cycle. Early electron microscope work showed that the calcified cell walls of coralline algae have a two-layered structure: An inner layer of acicular calcite parallel to the cell wall, succeeded by radial, inward growing calcite crystals (Bailey & Bisalputra 1970; Alexandersson 1974; 1977; Flajs 1977a; b; Garbary 1978; Cabioch & Giraud 1986; Bosence 1991). Microprobe plotting of magnesium and calcium concentrations by Flajs (1977b) and Massieux et al. (1983) indicated that the secondary radial layer has a higher magnesium content than the earlier parallel layer. Because of that, the Mg/Ca ratio should decrease towards the end of a growth period, where the cell rows are less calcified than at the beginning of the growth period. Testing this pattern against the Mg/Ca ratio using electron dispersive X-ray (EDX) spectroscopy results in the correlation of high Mg/Ca ratios with rows of heavily calcified cell walls

growth direction

annual growth increment 0.16 0.14 0.12 0.08 0.10 Mg/Ca ratio Mg/Ca 0.06 0.02 0.04 0.00

Fig. 7.27. Mg/Ca transect measured by EDX along a longitudinal section of a Lithothamnion glaciale protuberance showing a decrease of the Mg/Ca ratio towards the end of an annual growth increment; scale bar = 50 µm.

70 Results and low Mg/Ca ratios with rows of less calcified cell walls (Fig. 7.27), giving rise to an annual banding pattern. The assumption of an annual banding pattern is furtherly confirmed by the distribution of the reproductive conceptacles in the protuberances of L. glaciale. These conceptacles contain the spores of the plants and form anually (Jackson 2003). Longitudinal, fuchsine-stained sections of protuberances of L. glaciale clearly exhibit that the pattern of the conceptacle distribution parallels the light-dark banding pattern of the growth increments (Fig. 7.28). This light-dark banding pattern resembles the heavily and less calcified cell rows.

conceptacles summer winter annual growth increment

Fig. 7.28. Longitudinal section of a fuchsine-stained protuberance of L. glaciale from the Nord- kappbukta site. The annual growth increments are visible and are divided in bright summer and dark winter bands, note that the annual forming conceptacles coincide with the banding pattern; scale bar = 500 µm.

71 Results

7.9 The annual carbonate production by the rhodoliths

After sorting of the samples, only the monospecific rhodoliths consisting of Lithothamni- on glaciale (see Table 7.4 for the number of samples and the features of the particular sites) were chosen to examine the living rhodolith surfaces with respect to the number of protuberances per unit area as described above. This approach resulted in the median number of protuberances per square metre. The results were checked for significance using Levene’s test for homogeneity of variance based on means (One-way ANOVA) and are compiled in Table 7.5. After the measurements regarding the numer of protuberances per unit area were completed, five protuberances each of randomly selected rhodoliths were cut off, longitudinal measured, dried, and weighed as described above. The protuberances were then prepared for increment counting by resin embedding, longitudinal sectioning, wet polishing, and fuchsine staining. The fuchsine-stained sections of the protuberances revealed clear growth increments, and where those were absent, the annual forming conceptacles were successfully used as a substitute, since they paralleled the light-dark banding pattern of the growth increments (see Fig. 7.28). Because the succession of the conceptacles corresponds with the banding pattern of the growth increments, the assumption that the growth increments are annual is furtherly confirmed. After the annual growth increments were counted, the consequently outcoming age of each protuberance was plotted against the total weight of the particular protuberance. The results show a clear linear increase of weight with age. This linearity between age and weight was checked for correlation using the reduced major axis algorithm showing a high coefficient of determination and checked for significance using Levene’s test for homogeneity of variance based on means (One-way ANOVA). The results are compiled in Fig. 7.29. The hence calculated mean carbonate productions per protuberance per year (see Table 7.6) for each locality were checked for significance using Levene’s test for homogeneity of variance based on means (One-way ANOVA). The calculated mean values of each five protuberances per rhodolith were then multiplied with the median numbers of protuberances per square metre of the belonging site, which led to the total carbonate production per square metre per -2 -1 year [g CaCO3 m yr ] for each site (see Table 7.7). These results together with the possibly influencing constants depth, geographical latitude, and duration of the polar night, and the possibly influencing variables duration of the sea ice cover, annual mean temperature, and seawater calcite saturation (see Table 7.8) were checked for their coherence using the reduced major axis algorithm and checked for significance using Levene’s test for homogeneity of variance based on means (One-way ANOVA). The results from site 714 (Nordkappbukta) were excluded because of their strongly differing water depth, which adulterated the coherence checks. The results are compiled in Fig. 7.30 and while coherences for water depth and seawater calcite saturation are not evident, there is a clear coherence for the annual mean temperature and a very strong correlation with the geographical latitude, the duration of the polar night, and the duration of sea ice cover.

72 Results

Table 7.4. The rhodolith sampling sites in order from south to north together with the number of rhodoliths collected all in all and the number of rhodoliths being sufficient for further analysis (Lat. [°N] = geographical latitude in decimal degrees; Long. [°E] = geographical longitude in decimal degrees).

Locality Site # depth [m] Lat. [°N] Long. [°N] Rhodoliths collected / sufficient Floskjeret 757 45 78.3113 14.5328 37 / 17 Krossfjorden 652 47 79.0618 11.2291 2 / 2 Krossfjorden 637 50 79.0985 10.7940 104 / 31 Krossfjorden 644 41 79.0986 10.7603 15 / 15 Mosselbukta 684 44 79.8946 15.7428 77 / 27 Nordkappbukta 714 27 80.5251 19.8691 42 / 37 Nordkappbukta 711 45 80.5293 19.6673 43 / 28 Nordkappbukta 701 38 80.5335 19.8461 23 / 10

Table 7.5. Number of protuberances per square metre rhodolith surface with the median, minimum, and maximum values and Levene‘s test showing the levels of significance.

Protuberances / surface [n m-2] Locality Site # Samples [n] Median Min Max Levene’s test Floskjeret 757 17 35239 14454 55636 p < 0.01 Krossfjorden 652 2 55302 51248 55302 p < 0.001 Krossfjorden 637 31 71096 30446 141810 p < 0.001 Krossfjorden 644 15 59673 22903 105201 p < 0.05 Mosselbukta 684 27 28499 19380 60094 p < 0.001 Nordkappbukta 714 37 30740 9953 65088 p < 0.001 Nordkappbukta 711 28 21404 8946 46883 P < 0.01 Nordkappbukta 701 10 22179 11967 40903 p = 0.121

73 Results

100 100 90 90 80 80 70 70 60 60 50 50 40 40 30 757 30 652 2 20 R2 = 0.834 20 R = 0.875 p < 0.001 p < 0.001 annual growth increments / age [yrs] increments annual growth annual growth increments / age [yrs] increments annual growth 10 n = 60 10 n = 10 0 0 0 100 200 300 400 500 600 700 800 900 0 100 200 300 400 500 600 700 800 900 weight [mg] weight [mg] 100 100 90 90 80 80 70 70 60 60 50 50 40 40 30 637 30 644 2 20 R = 0.891 20 R2 = 0.636 p < 0.001 p < 0.001

annual growth increments / age [yrs] increments annual growth 10 n = 35 / age [yrs] increments annual growth 10 n = 30 0 0 0 100 200 300 400 500 600 700 800 900 0 100 200 300 400 500 600 700 800 900 weight [mg] weight [mg] 100 100 90 90 80 80 70 70 60 60 50 50 40 40 30 684 30 714 2 20 R2 = 0.819 20 R = 0.886 p < 0.001 p < 0.001

annual growth increments / age [yrs] increments annual growth 10 annual growth increments / age [yrs] increments annual growth 10 n = 60 n = 35 0 0 0 100 200 300 400 500 600 700 800 900 0 100 200 300 400 500 600 700 800 900 weight [mg] weight [mg] 100 100 90 90 80 80 70 70 60 60 50 50 40 40 30 711 30 701 2 20 R2 = 0.791 20 R = 0.831 p < 0.001 p < 0.001 annual growth increments / age [yrs] increments annual growth annual growth increments / age [yrs] increments annual growth 10 n = 35 10 n = 35 0 0 0 100 200 300 400 500 600 700 800 900 0 100 200 300 400 500 600 700 800 900 weight [mg] weight [mg]

Fig. 7.29. Scatter plots of the number of annual growth increments against the total weight of the particular protuberance for all sites. The plots indicate a linear weight increase with age and show a high coefficient of determination and a high significance indicating a strong correlation of the values.

74 Results

Table 7.6. Mean carbonate production per protuberance per year calculated from the annual growth increments in relation to protuberance weight with the median, minimum, and maximum values and Levene‘s test showing the levels of significance.

Weight / annual growth increment [mg n-2] Locality Site # Samples [n] Mean Min Max Levene’s test Floskjeret 757 60 5.68 3.00 11.18 p < 0.001 Krossfjorden 652 10 3.07 2.25 3.93 p < 0.01 Krossfjorden 637 35 2.55 1.43 4.21 p < 0.001 Krossfjorden 644 30 2.84 1.82 4.24 p < 0.001 Mosselbukta 684 60 4.20 2.17 8.41 p < 0.001 Nordkappbukta 714 35 5.13 2.59 9.64 p < 0.001 Nordkappbukta 711 35 4.92 2.54 8.32 P < 0.001 Nordkappbukta 701 35 4.55 1.55 8.57 p < 0.001

Table 7.7. Annual carbonate production per square metre per year at each site calculated from the mean weight per annual growth increment and the median number of protuberances per rhodolith surface.

Locality Site # Mean weight / band1 Median protuberances / surface Annual carbonate production [mg n-2] [n m-2] [g m-2 yr-1] Floskjeret 757 5.68 35239 200.3 Krossfjorden 652 3.07 55302 169.8 Krossfjorden 637 2.55 71096 181.5 Krossfjorden 644 2.84 59673 181.5 Mosselbukta 684 4.20 28499 119.8 Nordkappbukta 714 5.13 30740 157.7 Nordkappbukta 711 4.92 21404 105.3 Nordkappbukta 701 4.55 22179 100.9 1 meaning one annual growth increment.

Table 7.8. Annual carbonate production per square metre per year at each site together with the possibly influencing parameters (Lat. [°N] = geographical latitude in decimal degrees; CalΩ = calcite saturation).

1 2 3 Locality Site # Annual carbonate production Depth Lat. Polar night Sea ice T ΩCal [g m-2 yr-1] [m] [°N] [days] [months] [°C] Floskjeret 757 200.3 45 78.3113 113 4 1.5 1.44 Krossfjorden 652 169.8 47 79.0618 117 6 0.7 2.67 Krossfjorden 637 181.5 50 79.0985 118 6 0.7 1.45 Krossfjorden 644 181.5 41 79.0986 118 6 0.7 1.45 Mosselbukta 684 119.8 44 79.8946 122 7 0.7 2.59 Nordkappbukta 714 157.7 27 80.5251 126 10 0.5 2.86 Nordkappbukta 711 105.3 45 80.5293 126 10 0.5 2.86 Nordkappbukta 701 100.9 38 80.5335 126 10 0.5 2.86 1 data from USNO Sun Rise Tables. 2 data from Węslawski et al. 1995; Svendsen et al. 2002; Nilsen et al. 2008; Spreen et al. 2008; AMSR-E Sea Ice Maps. 3 data from LEVITUS 94.

75 Results

81.0 11 a R2 = 0.953 b R2 = 0.866 p < 0.001 p < 0.005 80.5 n = 7 n = 7 9 80.0

79.5 7

79.0 5 geographical latitude [°N] latitude geographical 78.5 duration of sea ice cover [months] cover of sea ice duration

78.0 3 90 110 130 150 170 190 210 90 110 130 150 170 190 210 annual carbonate production [g m-2 yr-1] annual carbonate production [g m-2 yr-1]

130 2.0 c R2 = 0.929 d R2 = 0.480 p < 0.001 p < 0.05 n = 7 n = 7 125 1.5

120 1.0

115 0.5 duration of polar night [days] of polar night duration annual mean temperature [°C] annual mean temperature

110 0 90 110 130 150 170 190 210 90 110 130 150 170 190 210 annual carbonate production [g m-2 yr-1] annual carbonate production [g m-2 yr-1]

55 3.0 e f R2 = 0.745 p < 0.05 n = 7 50 2.5

45 Cal 2.0 Ω depth [m]

40 1.5 R2 = 0.218 p < 1 n = 7 35 1 90 110 130 150 170 190 210 90 110 130 150 170 190 210 annual carbonate production [g m-2 yr-1] annual carbonate production [g m-2 yr-1]

757 (Floskjeret) 637 (Krossorden) 684 (Mosselbukta) 701 (Nordkappbukta) 652 (Krossorden) 644 (Krossorden) 711 (Nordkappbukta)

Fig. 7.30. Scatter plots of the annual carbonate production by the rhodolith beds against possibly influencing parameters. There are very clear coherences and high significances for (a) the geographical latitude, (b) the duration of sea ice cover, and (c) the duration of the polar night; a coherence for (d) the annual mean temperature is also evident and shows a high significance. There is no coherence for (e) the water depth and (f) the calcite saturation; note that the apparent correlation with the calcite saturation is not reasonable (see discussion for explanation).

76 Discussion

8. Discussion

8.1 Environmental controls on Arctic rhodoliths

The hydrodynamic regime

The morphology of non-nucleated rhodoliths directly reflects the hydrodynamic regime (Bosence 1983b; Steneck 1986). Ellipsoidal shapes are more easily transported than spheroidal forms, and discoidal rhodoliths are most resistant against displacement through water currents (Bosence 1983b). Extensive beds of rhodoliths and also of maërl are found in areas with moderate to strong seabed currents, which are relatively open but yet sheltered enough to inhibit physical destruction of the often fragile coralline algal skeletons (Bosence 1979). The shape of nucleated rhodoliths that bear lithoclasts is widely predetermined by the particular nuclei (Freiwald 1995). Nevertheless, the shape of the lithoclasts also depends on the hydrodynamic regime, even if on a larger scale of time (Sneed & Folk 1958), so the shapes prevailing in a particular rhodolith bed are also related to the hydrodynamic regime. It is obvious that these beds require both shelter from wave action to prevent burial and destruction of the thalli but also enough water movement to avoid smothering with fine sediment (Hall-Spencer 1998). While burial in coarse sediment has less severe effects on the algae, burial in fine sediment mostly leads to the dieback of the thalli due to the loss of light incidence as needed for photosynthesis (Wilson et al. 2004). The rhodolith beds from the four investigated sites along the coast of Svalbard are also exposed to a variety of hydrodynamics and hence show different responses.

Floskjeret in the outer part of Isfjorden is regularly exposed to the influences of the ESC and the WSC as well, which results in regularly occurring hydrographic switches from ArW (transported by the ESC) to AW (transported by the WSC) conditions and vice versa (Cot- tier et al. 2007; Nilsen et al. 2008). Because of that, the site is indeed subject to the impact of strong currents, but since the switches occur only twice a year (Nilsen et al. 2008), they have no distinctive impact on the shape of the prevailing lithoclasts. Since Floskjeret is situated in the sheltered bay of Borebukta (Fig. 2.4b), it rather appears to be a site that offers protection from strong waves and currents. These conditions mirror in the morphology of the prevailing rhodoliths and their lithoclastic nuclei. Since there is no concentration on a particular shape (43% discoidal, 38% spheroidal, 19% ellipsoidal; Fig. 7.22a), the hydrodynamic impact seems to be low, not favouring the development of spheroidal forms (Sneed & Folk 1958). This also coincides with the large size range of the rhodoliths (2.9-8.1 cm; Fig. 7.23a), while a higher hydrodynamic energy would result in smaller lithoclastic nuclei (Bloore 1977). Also the well-developed, large protuberances up to >60 years old (Fig. 7.29; station 757), which are not subject to distinct abrasion, argue for such conditions.

77 Discussion

Krossfjorden is strongly affected by currents due to the interaction between the ESC and the WSC (Svendsen et al. 2002). The examined rhodolith communities prevail predominantly at the shallow Mitragrunnen Bank (Fig. 2.4c). This bank is offshore from the mouth of Kross- fjorden, so the hydrodynamics exerted by the currents are further increased by wave action (Svendsen et al. 2002). This high energy environment leads to predominantly spheroidal (58%) and ellipsoidal (29%) shapes of the colonised lithoclasts (Fig. 7.22b) and a small size range (1.6-2.7 cm; Fig. 7.23b) due to intense abrasion processes (Sneed & Folk 1958; Bloore 1977). It also coincides with the observation that the rhodoliths from Krossfjorden are at a low state of development, mostly consisting of lithoclasts thinly encrusted by coralline algae with poorly matured protuberances. Another reason for this low state of rhodolith development may be the frequent ice run at the Mitragrunnen Bank. The JAGO underwater video footage shows many iceberg scouring- marks indicating that this is a common feature at Krossfjorden and especially at Mitragrun- nen Bank due to its shallow water depth. These observations are confirmed by the findings of Svendsen et al. (2002), reporting on numerous iceberg drifts in that area. This suggests that the rhodolith communities are regularly interrupted in their maturing process by iceberg scouring and consequently are poorly developed, consisting only of small specimens with thin crusts and often not well-developed protuberances, even in water depths were the light regime would offer sufficient conditions. Additionaly to the limitations exerted by the hydrodynamic conditions, another reason for the low level of rhodolith develoment might be that the Krossfjorden community is still young compared with the other examined rhodolith beds. This is also likely regarding the counting of the protuberance increments, where the oldest examined protuberance from Krossfjorden exhibited an age of only 42 years (Fig. 7.29; station 652), while all other sites yielded ages of more than 60 years (Fig. 7.29).

The area of Mosselbukta and Wijdefjorden is exposed to the impacts of both the ESC and the WSC, and a mixing of AW and ArW occurs. The impact of AW is only pronounced during a period of strong activity of the WSC; during the rest of the time, the influence of ArW prevails (Sapota et al. 2009). Hence, the site is subject to the impact of strong currents only in few times of changing conditions (Sapota et al. 2009), so they have no distinctive impact on the shape of the prevailing lithoclasts. Since Mosselbukta is a sheltered bay (Fig. 2.4d), it rather appears to be a site that offers protection from strong waves and currents. The reason why the shapes of the prevailing lithoclasts and hence of the rhodoliths concentrate on spheroidal forms (79%; Fig. 7.22c) might be that the hydrodynamic regime exerted a stronger force in times before the rhodolith beds developed. This would have led to spheroidal lithoclasts (Sneed & Folk 1958; Bloore 1977) that were later colonised by coralline algal spores as described by Freiwald (1995). The examined rhodoliths also exhibit a large range of size (1.9-8.8 cm; Fig. 7.23c) which suggests an advanced age of the rhodolith communities. Supporting this assumption, the oldest protuberance from Mosselbukta used for increment counting exhibited an age of >70 years (Fig. 7.29; station 684).

78 Discussion

Nordkappbukta is located at the northern tip of the Laponiahalvøya, Nordaustlandet, where it is exposed to influences of the ESC and the WSC, but the impact of the AW is only pronounced in periods of strong WSC activity, whereas at other times the area is under the influence of ArW (Sapota et al. 2009). These changes result in strong, but only seasonally occurring currents (Sapota et al. 2009), that likely do not have a pronounced impact on the lithoclastic material and hence not on the shapes of the rhodoliths. The reason why the shape of the prevailing lithoclasts and hence of the rhodoliths concentrate on spheroidal forms (60%; Fig. 7.22d) might be the same as at Mosselbukta, namely that the hydrodynamic regime exerted a stronger force in times before the rhodolith beds developed, which would have led to spheroidal lithoclasts (Sneed & Folk 1958; Bloore 1977). The examined rhodoliths also exhibit a large range of size (1.8-8.4 cm; Fig. 7.23d) which suggests an advanced age of the rhodolith communities. The observation that the oldest protuberance from Nordkappbukta used for increment counting exhibited an age of >90 years (Fig. 7.29; station 701) supports this assumption. Additionally, the site is also affected by iceberg scouring, but as this feature was observed less frequent during the JAGO dives than at Krossfjorden, it does not seem to have a pronounced impact on the development of the rhodolith communities.

Taken as a whole, the shapes of the rhodoliths from the four sites cannot be used as indicators for the prevailing hydrodynamic regime. Since all rhodoliths are nucleated, or, in the case of the hollow rhodoliths (Fig. 7.26), presumably contained a nucleus at an earlier growth stage, their shape is widely predetermined by their particular nucleus as stated by Freiwald (1995). The lithoclastic nuclei, that are encrusted by coralline algae, however, may be used as indicators for former hydrodynamic conditions (Sneed & Folk 1958; Bloore 1977). In this coherence, if the predominant rhodolith shapes and size ranges do not coincide with the prevailing hydrodynamic conditions, as it holds for Mosselbukta and Nordkappbukta, the onset of conditions favourable to rhodoliths could be approximately determined by the age of the rhodolith communities. At that scale, the specimens from Svalbard confirm the importance of rhodolith beds as potential indicators for palaeoenvironmental conditions (Foster 2001).

The substrate

Even if light is the most limiting factor to the coralline algae (Kain & Norton 1990), their abundance only increases with decreasing water depth if adequate substrate is available. The substrate has to offer a distinct stability and should be free from fine sediment that decreases the photosynthetic efficiency of the coralline algae (Wilson et al. 2004). In Svalbard, these dependencies result in a patchy distribution of the rhodolith beds and hence in several spatially separated communities. Most commonly, the coralline algae are attached to lithoclasts, and even if corallines can also occur as epiphytes growing on the surface of other organisms (Kain & Norton 1990), shell accumulations (mainly consisting of C. islandica and H. arctica) are of much lesser importance. The light shells, as well as cobbles below

79 Discussion a distinctive size, do not provide enough stability, so stormwaves and strong currents would intensely affect the coralline algal crusts (Adey 1970a). Bare bedrock was only observed at the Nordkappbukta site (JAGO dive track 701; see also Fig. 7.9b), and was widely covered with well-developed coralline algae, but presumably glaciogenic debris were commonly observed at all dive tracks. Under suitable conditions of water depth and water movement, these cobbles were colonised and rhodoliths developed. Apart from providing enough stability, the colonised substratum also has to be free from fine sediment as it regularly occurs at the Svalbard sites. Hence, the hydrodynamic regime acts as an important part as e.g. at Nordkappbukta, where the prevailing currents keep the exposed surfaces free from fine sediments (Sapota et al. 2009). Where rhodolith beds prevail, the topography of the seafloor is mostly flat, so the fine sediment deposition is limited to depressions, and these depressions are generally devoid of living rhodoliths. The reason for this phenomenon is that the fine sediment strongly hinders the photosynthesis of the coralline algae, thus leading to a successive dieback of the organisms (Wilson et al. 2004), while burial with coarse sediments, that cause a minor decrease of the PAR, has much less severe effects on the algae. The sedimentation of the fine sediments is probably of seasonal origin due to melt water transport from the glaciers, which bear an immense sediment load (Forwick & Vorren 2009). The initial formation of coralline algal crusts at Svalbard usually starts on presumably glaciogenic lithoclasts consisting of moraine gravel and dropstones. Larger cobbles and boulders >1 m3 are colonized preferentially because they offer the best stability and light exposure. Layers of fine sediments on lithoclasts hinder the colonisation and sediment-trapping depressions are always free from rhodoliths for the reasons mentioned above. The successive development of the coralline algae from an initial stadium of encrustation to well-developed rhodoliths is clearly visible along single dive tracks (Fig. 8.1) and follows the same pattern. From the depth where irradiance is sufficient (c. 78, m averaged for all stations), boulders >1 m3 and the larger lithoclast are colonised by initial crusts of coralline algae (Fig. 8.1b). With decreasing water depth and the coeval increase of PAR, the crusts begin with the development of small protuberances. These protuberances grow observably longer and thicker and become increasingly denser on the coralline surface (Fig. 8.1c). This development continues until the coralline crust constitutes >50% of the nodule consisting of the lithoclast and coralline algal skeleton, henceforward it is called rhodolith per definition (Adey & Macintyre 1973; Bosence 1983a; b). Regarding this, the development of rhodolith colonisation follows a multistep process and is progressed at different rates at the Svalbard sites. In Krossfjorden, the development of rhodolith beds is at an early state and exhibits thin, smooth crusts on presumably glaciogenic debris. With further development, the thickness of the crusts increases and the first small protuberances appear, marking the changeover from basic coralline algal crusts to rhodoliths with protuberances of up to 42 years in age (Fig. 7.29; station 652). In the next step, as it is exhibited in Floskjeret, the rhodolith forming coralline algal crusts are already several millimetres thick and the protuberances are numerous and well developed, with ages of partly >60 years (Fig. 7.29; station 757). Lastly, the rhodoliths from Mosselbukta and Nordkappbukta show

Discussion

depth depth 30 m 80 m 30 m 70 m a d 500 m 250 m

. 2012). coralline algal-cover and -development and algal-cover coralline distance bedrock

gravel and shells on silt gravel coralline algal-cover and -development and algal-cover coralline bedrock ( c ) bedrock 0 m distance gravel and shells on silt gravel gravel and boulders on silt ( f ) gravel gravel on silt ( e ) gravel DIVE TRACK DIVE TRACK 701 gravel and boulders on silt gravel gravel and shells on silt gravel „Polysiphonia-“ fouling „Polysiphonia-“ iceberg plough mark iceberg with bleached corallines boulders on silt ( b ) gravel on silt gravel gravel encrusted by encrusted gravel algae coralline by boulder encrusted algae coralline gravel algal coralline crusts on bedrock shell bed 0 m gravel, boulders and shells on silt gravel, gravel and shells on silt gravel c DIVE TRACK DIVE TRACK 711 gravel, boulders and shells on silt gravel, gravel and shells on silt gravel f b e Facies profiles and associated seafloor pictures of JAGO dive tracks 701 and 711 showing the predominant pattern of coralline algal settlement and rhodoltih and coralline algal of settlement pattern predominant the showing 711 and 701 tracks dive JAGO of pictures associated seafloor and profiles Facies 8.1. Fig. to algal coralline crusts attached ( c ) well-developed depth; algal initial coralline with crusts 701; ( b ) boulders in 75 m water track of dive ( a ) profile development; ( f ) bleached depth; in 45 m water bottom a gravel on development of stage intermediate an at 711; ( e ) rhodoliths track of dive ( d ) profile depth; bedrock in 51 m water et al Teichert after modified = 10 cm (slightly scale depth; in seafloor images bars in 36 m water algal scouring-mark iceberg red crusts in an

81 Discussion even larger and denser protuberances than those as exhibited in Floskjeret, some protuberances with ages >70 years in Mosselbukta (Fig. 7.29; station 684) and >90 years in Nord- kappbukta (Fig. 7.29; station 701). Additionally, many of the rhodoliths from Mosselbukta and Nordkappbukta seem to have lost their lithoclastic nucleus resulting in hollow structures. The mechanism of the formation of hollow rhodoliths is not fully understood, but it seems that the rhodoliths from Svalbard always start as initial coralline algal crusts and lose their lithoclastic nucleus at some state of their development (see Fig. 7.17d). After that, they presumably grow further at their meristematic rims, successively closing the gaps between the several parts of the crusts, finally forming a hollow, more or less spheroidal shaped structure

The great dependence of coralline algae that form rhodoliths on an appropriate substrate also implies the possible outcome of increasing temperatures due to the ongoing global change. On the one hand, glaciers that now transport large cobbles and boulders, being an excellent substrate for the settlement of coralline algae, may lack this capacity if their volume, mass and the associated transport mechanisms decrease (Dowdeswell et al. 1997), so the supply of appropriate substrate will peak off. At the same time, the progressively melting glaciers will transport a much bigger load of fine sediment to the seafloor, thus resulting in impaired conditions for the photosynthesis of the coralline algae (Wilson et al. 2004). Such changes may most likely result in the successive dieback of the rhodolith communities. Global change models also predict changes in coastal currents (Cottier et al. 2005) and increasing wind forces for the Arctic regions (Hinzman et al. 2005), that would lead to a modified hydrodynamic regime. If the the water energy increases, lithoclasts formerly stable enough to enable the settlement and development of coralline algal crusts might be transported towards regions with unfavourable conditions for the algae. Regarding the limited availability of substrate, another problem that might arise with the global change is the invasion of new species (Cheung et al. 2009). If temperatures further increase, species with Atlantic origin will migrate northwards, a trend that is already observable from the species composition during this study, as it e.g. holds for the Atlantic snake pipefish (Entelurus aequoreus), a species that was presumed to be confined to the area south of Iceland (Fleischer et al. 2007). This migration might not be limited to vagil, but also to fixosessil benthic organisms, which then act as competitors for space to the coralline algae. Another scenario, as suggested by Konar & Iken (2005), is that the climate change may also influence benthic communities through an alteration of the disturbance systems. As there is a significant loss of Arctic sea ice predicted to occur by 2025 (Clarke & Harris 2003) especially in coastal regions (Morison et al. 2000), Konar & Iken (2005) suggest that Arctic nearshore systems would actually experience more disturbances as a result of the climate change. Since a lack of shore fast-ice cover could result in an increase in wave action in such shallow waters, the impact on the residing rhodolith communities would be substantial.

82 Discussion

Temperature

Temperature has great effects on respiration, photosynthesis and growth rates of coralline algae (Kain & Norton 1990; Wilson et al. 2004). The optimum temperature clearly varies geographically and with species, but the general pattern usually shows an increase in growth rate to a maximum that is near the top of the tolerated range (Kain & Norton 1990). Hence, temperature is the primary determinant of geographical species distribution, and the boundaries of biogeographical regions are associated with isotherms (Lüning 1990). Parallel to this, Adey & Adey (1973) showed that the distribution patterns of coralline algae may be corre- lated with temperature boundaries. The maximum coverages at the different sites at Svalbard appear at c. 45 m water depth and the potential mean temperatures in that water depth vary from 0.5°C at Nordkappbukta to 1.5°C at Floskjeret (data from LEVITUS 94). These relatively high temperatures result from the intermixing of the warm AW deriving from the WSC with the colder ArW deriving from the ESC (Orvik & Niiler 2002; Sapota et al. 2009). However, the temperatures are low compared to e.g. boreal localities (Lüning 1990), and its common appearance at the Svalbard sites shows that L. glaciale is adapted to these low temperatures compared to e.g. Lithothamnion corallioides (Crouan & Crouan) Crouan & Crouan, 1867, which has a minimum survival temperature of 5°C (Adey & McKibbin 1970). On the other hand, it is also temperature that seems to limit the southward distribution of L. glaciale, possibly because the reproductive conceptacles are only produced when the water temperatures are <9°C (Hall-Spencer 1994). The CTD records (Fig. 7.11) only show snapshot states but they were taken during summer time and under ice-free conditions. Hence, one can assume that they may show values close to the possible maxima. The water temperatures in c. 45 m water depth, where the coverage with and the development of the rhodoliths reach their maxima, are far below the 9°C limit, so the formation of conceptacles is presumably possible throughout summertime. This is a great advantage, since the reproductive frequency of L. glaciale is annually protracted (Jackson 2003), and reproduction during summer implies sufficient light conditions, while it may fail during winter darkness. Altogether, L. glaciale occupies a distinctive temperature range at Svalbard, which is coequally warm enough to enable sufficient growth and cold enough to enable reproduction during summer.

According to the remarks above, it becomes clear that the coralline algae prevailing at the four sites depend on a distinct window in the temperature range. If the winter mean temperatures would decrease, the vital processes may be disrupted (Kain & Norton 1990), which would lead to a deferred start at the beginning of the growing season, or even to a complete dieback of the thalli. On the other hand, strongly increasing mean temperatures would possibly hinder the formation of reproductive conceptacles during the growing season (Hall-Spencer 1994), so the settling gametes were not able to leverage the enhanced conditions of the light regime and hence would go under other, faster growing plants.

83 Discussion

Salinity

Fluctuations in salinity may cause osmotic stress, unfavourable ionic balances and a shortage of essential metabolites (Kain & Norton 1990). Passing down a salinity gradient, the number of species of the Rhodophyta commonly declines sooner than that of the Phaeophyta, whereas that of the Chlorophyta may actually increase (Coutinho & Seeliger 1984; Kain & Norton 1990). The oceanographic regimes of the Svalbard sites are strongly influenced by the interactions between the ESC and the WSC, and the CTD records indicate a stratified summer situation, characterised by warm surface AW from the WSC overlaying a mixing zone with colder ArW from the ESC. The salinity measurements show a thin incumbent layer of less saline water, being a result of the strong freshwater input from the melting glaciers, such as Lilliehööbreen in the Krossfjorden transect (Fig. 7.12a). Because of the lower density of the less saline melt water layer, it does not fully intermix with the water masses from the ESC and the WSC, which feature fully marine conditions (Svendsen et al. 2002). Thus, the conditions in the water depths where well-developed rhodoliths occur (c. 45 m) remain fully marine even in times with increased melt water input. Nevertheless, these apparently stable conditions are only a snapshot and may change during times of melting sea ice or pronounced shifts of ocean currents and coherent mixtures of AW and ArW (Svendsen et al. 2002).

However, the prevailing conditions are suitable to L. glaciale, which favours fully marine conditions but is also able to tolerate both low (<18, Wilson et al. 2004) and strongly fluctuating salinities (18-40 in sea lochs off the west coast of Scotland, Jackson 2003). Therefore, the seasonal decreasing salinities induced by melting glaciers during summer time likely have non or at most low direct impact on the rhodolith communities. The adaptation to varying salinities is an important attribute of L. glaciale that contributes to its persistence in various environments. It may also partly explain its predominance towards P. tenue, that is not known to possess high salinity tolerances. Though, it should be stressed that L. glaciale indeed tolerates low and fluctuating salinities but is best adapted to fully marine conditions. This becomes also evident from the experiments done by Adey (1970b) and Wilson et al. (2004: 283), showing that L. glaciale survives even very low salinities but with strongly decreased photosynthetic capacities. This implies that permanently decreased salinities also in deeper water layers, as it may be the consequence of a strongly increased glacier meltdown (Svendsen et al. 2002; Clarke & Harris 2003), would negatively affect the rhodolith beds and lead to impaired growth conditions.

Irradiance

Coralline and other red algae can tolerate a wider range of light levels than any other group of photosynthetic plants, and many are low-light-adapted (Kain & Norton 1990). In over 90% of the red algal species, chlorophyll a and several carotenoids occur along with the accessory

84 Discussion pigments phycoerythrin and phycocyanin (Ragan 1981), so their absorption spectrum is much higher than that of the green algae, which only contain chlorophyll a, chlorophyll b, and several carotenoids (Fig. 3.1). This holds both for geniculate (jointed) corallines (Hader et al. 1996) and non-geniculate corallines (Kühl et al. 2001; Roberts et al. 2002), and such species often show an effective adaptation to low irradiances in polar latitudes, which are often prolonged under sea ice conditions. Lüder et al. (2002) reported that the non-calcareous red alga Palmaria decipiens (Reinsch) Ricker, 1987 could cope with complete darkness for several months before respiration suddenly drops, while the photosynthetic capacity recovers rapidly after exposure to illumination, and Adey (1970b) stated that coralline algae of the high Arctic may require only one month of photosynthesis at low light levels for three fourths of each day to be able to live in the dark for the remainder of the year. This enables sufficient growth even at high latitudes like in Svalbard, where the polar night lasts for up to 126 days in Nordkappbukta (data from USNO Sun Rise Tables) and thus, L. glaciale and P. tenue seem to be adapted very well to long-term dark periods. Appearance and properties of the rhodolith beds slightly vary along the different dive tracks, but show the same overall pattern. The initial growth of encrusting coralline algae starts at c. 78 m water depth in the dysphotic zone (0.01-1% surface illumination), where the measured PAR is 0.1 µmol photons m-2 s-1 on average (Fig. 7.13). Roberts et al. (2002) showed that individuals of P. tenue have no significant net photosynthesis at such low irradiance. At 45 m water depth (2.1 µmol photons m-2 s-1, averaged for all stations), rhodoliths cover nearly 100% of the hard substrate and the protuberances of L. glaciale are well-developed. The average abundance of P. tenue is 10% at Nordkappbukta, and data from Roberts et al. (2002) show a net photosyn- -2 -1 thesis of only 2 mmol O2 m d for individuals of this species at such irradiance conditions. Hence, the initial appearance of L. glaciale at >75 m water depth shows that these algae can cope with very low light conditions and are able to colonise a large range of the dysphotic zone. It might also be one reason why there is such disequilibrium between the abundance of L. glaciale and P. tenue. Wilson et al. (2004) showed that coralline algae might lack the ability to perform additional photochemistry under high irradiance conditions. The decrease in coverage with further to- pographic flattening (<30 m water depth) seems to be directly linked to the increasing coverage of Polysiphonia-like red algae, which do not calcify, grow much faster than the coralline algae and find sufficient light conditions in the shallow waters. However, this seaweed cover seems not to be dense enough to enable the development of a shaded coralline algal understo- rey at very shallow depths (see Irving et al. 2005).

Irradiance is another factor that limits the appearance of coralline algae at Svalbard to a distinct depth gradient, since light conditions seem to be insufficient in water depths >78 m, while in a depth <30 m, irradiance is high enough to favour other plants, which displace the coralline algae. The nearly 50-m-wide depth range for L. glaciale is much larger than characteristic depth ranges for algae in temperate and tropical environments. This observation indicates a very high degree of shade adaptation as it has been demonstrated for coralline algae

85 Discussion from the Ross Sea, Antarctica (Schwarz et al. 2005). The differences in low-light-adaptation also seems to be one reason for the disequilibrium between the amount of L. glaciale and P. tenue thalli. Lithothamnion glaciale can both occupy a much wider depth gradient and, since it recovers very fast from periods of complete winter darkness, it has an advantage at the beginning of the growing season (Wilson et al. 2004). Regarding that, the outcome of a potential global warming for the light regime and hence the thriving of the coralline algae at Svalbard is ambiguous. On the one hand, increasing temperatures would lead to shorter periods of sea ice cover and hence, since sea ice cover also spreads over the time period of the polar day (data from AMSR-E Sea Ice Maps and USNO Sun Rise Tales), strongly prolong the growing season of the coralline algae. On the other hand, this would also affect faster growing plants without low-light-adaption, so the coralline algae would lack the advantage during the times of the formation and the breakup of the sea ice, at which the light conditions are presumably too low for the growth of the Chlorophyta, hence giving the Rhodophyta a head start. A major problem arising with increasing annual mean temperatures is the simultaneous melting of the glaciers, resulting in the transport of huge amounts of fine sediments to the water column (Konar & Iken 2005). This would strongly impair the growing conditions for the coralline algae due to a reduced PAR and the parallel burial with the fine sediment, being a major reason for the dieback of coralline algae (Wilson et al. 2004).

Carbonate saturation

Because coralline algae are heavily calcified, and calcification involves high magnesium calcite precipitation (Bosence 1991), the concentration of calcium in the water column is critical for the coralline algae (King & Schramm 1982), and the maintenance of potassium in algal cells, that is transported by the ion pumps in the cell membrane, depends on the presence of adequate quantities of calcium ions (Kain & Norton 1990). The seawater carbon content, associated with its pH, has a marked effect on the photosynthesis at the low salinities (Kain & Norton 1990) that can occur particularly during times of intense melt water input, and hence carbonate saturation is an important factor for coralline algal growth at high latitudes. Martin et al. (2008) showed that an increasing acidification of the seawater leads to a significant reduction in coralline algal cover, indicating that a lowered pH and the reduction of carbonate saturation are important factors that may affect the rhodolith beds.

The water chemistry measured at all sites (Table 7.1) shows carbonate saturation (ΩCal and

ΩArg ≥ 1) and a pH ≥ 7.85 for the whole area, which is an important factor for the thriving of the rhodolith beds. The only stations slightly undersaturated regarding aragonite were sampled in water depths of 417 m (ΩArg = 0.91) and 332 m (ΩArg = 0.92), and hence are negligible with respect to the growth conditions of the coralline algae. Additionally, they are still saturated regarding calcite.

86 Discussion

This is remarkable, because high latitude oceans should be the first to become undersaturated with respect to calcite and aragonite (Orr et al. 2005). Hence, it is one of the main depen-dences that may be affected by the ongoing ocean acidification resulting in impaired conditions for the growth of coralline algae at high latitudes. Modelling studies projected annual mean carbonate undersaturation as early as 2032 for the Arctic surface ocean if anthropogenic CO2 emissions follow the IPCC (Intergovernmental Panel on Climate Change) business as usual scenario (SRES A2; Steinacher et al. 2009). This also implies that a possible acidification could lead to a decrease in rhodolith abundance, since it has been shown that coralline algae are very sensitive to risen CO2 conditions and the coequally lowered pH, re- acting with a strong decrease in the percentage of area covered (Hall-Spencer et al. 2008). On the other hand, Alexandersson (1974; 1977) and Okazaki et al. (1982) state that coralline algae induce a microenvironment suitable for carbonate precipitation by metabolic excretion of alginic acid, so that the particular levels of carbonate saturation of the ambient water column is of minor importance for the calcification process. But this may only hold for as long as the water column is still at least slightly saturated with respect to calcite (ΩCal ≥ 1), so the algae can deploy the free calcium ions (King & Schramm 1982).

Nutrient and carbon depletions

Although not explicitly analysed in this study, the ecophysiological adaptation of the coralline algae to overcome the summer depletion of macro-nutrients (nitrogen, phosphorus) deserves some consideration. As stated before, these macro-nutrients are available during the dark winter period in the waters around Svalbard and thus can be utilised by the coralline algae instantaneously. Such an ecophysiological adaptation has been experimentally proven for some polar phaeophytes (see Wiencke et al. 2007; and further references therein). The carbon needed to maintain the metabolism, biomass and even growth during the period of the polar night and sea ice cover derives from carbohydrates as storage products. Such car- bohydrate products occur as starch grains (amylopectin) in the coralline algae (Turvey 1978; Freiwald 1993; Freiwald & Henrich 1994), which are formed photosynthetically during the illuminated period and deposited within the vegetative cell compartments. During the dark period, the carbohydrates can be remobilised and may act as carbon source (Freiwald & Henrich 1994). A similar pathway has been detected by Lüning et al. (1973) for some polar laminarian phaeophytes. During this study, many SEM images were taken in order to determinate the taxonomy of the coralline algae prevailing at the different sites. Some of these images also show the starch grains mentioned above in thalli of L. glaciale (Fig. 7.20) and hence indicate that the algae are well adapted to the demands of the polar night. The starch grains were not detected in SEM images of thalli of P. tenue, but testing the thalli with iodine indicates the presence of starch reservoirs due to the blue staining of the thalli (pers. obs.).

87 Discussion

8.2 Interactions between rhodoliths and other benthic organisms

Distribution, composition, and abundance of the benthic organisms (excluding rhodophy- ceans) prevailing at the study areas seem to be mainly controlled by light penetration and the kinetic energy regime (waves, currents and tides). As these factors depend very much on the water depth, a depth zonation is the most pronounced pattern in the distribution of the benthic assemblages (pers. obs.). Due to the influence of the warmer AW, which is transported by the WSC, most benthic species are of Atlantic origin and do also occur at boreal latitudes, beeing not adapted to fully Arctic conditions. Many approaches have shown that coralline algae mostly are in a very close connection with the benthic organisms prevailing in their environment (Milliken & Steneck 1981; Steneck 1983; 1986). This holds for both coralline algae which grow as crusts and those that form rhodoliths. The biofilms covering the surfaces of the rhodoliths represent a major source of food for many organisms like gastropods, crustaceans, and sea urchins (Kain & Norton 1990). At the same time, this grazing activity exerted by the benthic organisms is beneficial to the rhodoliths, because it keeps them free from epiphytes (Adey & Macintyre 1973) and is thus essential for their general thriving. Many calcareous crustose species seem to be dependent in some cir- cumstances upon browsing animals to remove epiphytes or competitors that might otherwise swamp the algae (Brawley & Adey 1981; Steneck 1982). Herbivory is often identified as the source of disturbance that keeps rhodoliths clean and healthy (Steneck 1983; 1986), while the most effective physical defence against intensive grazing is seen in the calcareous thalli of the Corallinaceae, which are much tougher than those of most algae (Steneck 1983; Littler et al. 1983; Watson & Norton 1985). Such calcareous forms are among the most grazer-resistant algae, although even these are not immune (Clokie & Norton 1974; Adey & Vassar 1975; Steneck 1983; Padilla 1984). Their adaptations to withstand the grazing are numerous and comprise characteristics like meristems protected by thick epithallial tissue, sunken conceptacles sheltered from low-impact grazing as exerted by chitons, and intensely branched morphologies providing protection from high-impact grazing as exerted by echinoids (Steneck 1983). The rhodolith forming coralline algae prevailing at the Svalbard sites, L. glaciale and P. tenue, are among these grazer-resistant calcareous forms, and prominent grazers like Tonicella rubra and Strongylocentrotus sp. are very common in the rhodolith beds, feeding on the biofilms mainly consisting of diatoms which are present on the rhodoliths (Fig. 7.21). Tonicella rubra and other molluscs (Puncturella noachina and Tectura sp.), which are feeding on the rhodolith surfaces, exert a relatively low grazing pressure, while the impact of the echinoids Strongylocentrotus sp. and the more infrequent Strongylocentrotus pallidus is higher due to the larger and more effective Aristoteles’ lantern. The higher frequency of the intensely branched L. glaciale compared to the relatively smooth surfaced P. tenue at the investigated sites may be partially caused by the high abundance of Strongylocentrotus sp., because ecological studies have shown that branches in some non-geniculate corallines are an effective defence against deep-grazing sea urchins, because they do not reach the algal tissue between the protuberances but are restricted to whose apices (Milliken & Steneck 1981; Steneck 1983).

88 Discussion

Competition for space is also important especially for encrusting organisms and many coralline algae monopolize or virtually occlude the substrate by abutting with neighbours to form a continuous sheet (Littler & Kauker 1984; Johnson & Mann 1986). This is also visible at the four study sites, where many pebbles and cobbles are encrusted as a whole and appear completely pinkish red. Here, coralline algal crusts, primarily of the same species, L. glaciale, compete against each other, resulting in white meristematic margins that raise between individual crusts, if there is no more free space left on a particular lithoclast (Fig. 8.2). The coralline algae depend on the sufficient substrate for settlement and since this substrate is limited, competition for space is a common feature at the Svalbard sites. Addition- ally, the coralline algae also have to compete with e.g. serpulids, barnacles, and bryozoans. Most of the well-developed rhodoliths and of course the initial coralline algal crusts contain a lithoclastic core. Secondary to that, completely hollow rhodoliths are common at Mosselbukta and Nordkappbukta and act as kind of microenvironment for the benthic animals. The mechanism of the formation of hollow rhodoliths is not fully understood, but it seems that the rhodoliths from Svalbard always start as initial coralline algal crusts and lose their lithoclastic nucleus at some state of their development (see Fig. 7.17d). After that, they presumably grow further at their meristematic rims, successively closing the gaps between the several parts of the crusts, finally forming a hollow, more or less spheroidal shaped structure. These hollow rhodoliths are very often inhabited by various organisms of different groups (see Figs 7.17; 7.26), comprising bivalves (mainly consisting of Chlamys islandica and Hiatella arctica), ophiurids, serpulids, and others. Since most of the animals were still living at the

a b

Fig. 8.2. Lithoclasts encrusted by L. glaciale from Krossfjorden at 50 m water depth; a( ) initial state of encrustation with free space on the lithoclast; (b) fully covered lithoclast, note raised meristematic rims at the borders between different coralline algal specimens; scale bars = 5 mm.

89 Discussion time of sample collection (Fig. 7.17a), it is obvious that the animals really capitalise the hollow rhodoliths as microenvironments. The presumably glaciogenic gravel flats at the four sites are quite bare of protective cavities, so the hollow rhodolith structures, beside the interstices between several rhodoliths, allocate valuable shelter to many organisms. Regarding this interaction, the rhodoliths act as ecosystem engineers (Nelson 2009), providing a highly diverse habitat on the otherwise unprotected glaciogenic flats, presumably even favouring the reproduction of many organisms. Similarly, other studies report on rhodolith beds as refugia for scallops (Kamenos et al. 2004a) and as a habitat for juvenile cod (Kamenos et al. 2004b).

The observations from the Svalbard sites clearly indicate the important role of the rhodolith beds for the whole prevailing ecosystem. The biofilms present on the surface of the rhodoliths represents a major source of food for many grazing organisms and coequally, the rhodoliths benefit from the cleaning process exerted by these organisms. The rhodoliths apparently act as ecosystem engineers and hence occupy a major role in the production of manifold ecological niches. A regression in the rhodolith stock due to a dieback of the coralline algae would presumably result in more unfavourable conditions for many benthic organisms due to the loss of habitat and food resources.

90 Discussion

8.3 Rhodolith carbonate production

Different approaches have been applied to coralline algal growth-rate measurements and calculations of their carbonate production (Bosence & Wilson 2003), such as in situ growth experiments on living thalli (Adey & McKibbin 1970; Payri 1997; Steller et al. 2007), counting presumed annual growth increments in relation to protuberance weight (Bosence 1980; Freiwald 1993; Freiwald & Henrich 1994), using Mg/Ca ratio relative to growth increments (Halfar et al. 2000; Schäfer et al. 2011), alizarin- (Rivera et al. 2004; Foster et al. 2007) or calcofluor white-staining (Maytone 2010), and high-resolution analysis of 14C values to identify pre- and post-bomb spike periods of skeletal growth (Frantz et al. 2000). These completely different approaches, the use of not standardised sampling methods, as well as the handling of miscellaneous species of rhodolith forming coralline algae deriving from different water depths and biogeographical zones (see Table 8.1) make it difficult or nearly impossible to compare the resulting annual rhodolith carbonate production rates. In this study, the the carbonate production rates by the rhodolith beds, consisting mainly of L. glaciale, are based on annual increment counting of fuchsine stained protuberances in combination with the median number of rhodolith protuberances per square metre. The annual pattern of the growth increments is supported by the EDX-measured Mg/Ca ratio and the concomitant appearence of the annual forming conceptacles (Jackson 2003).

Table 8.1. Compiled annual carbonate productions per square metre per year by coralline algae.

Locality Lat. Depth Species carbonate production Reference -2 -1 [°] [m] [g CaCO3 m yr ] Floskjeret, Svalbard 78.31°N 45 Lithothamnion glaciale 200.3 this study Krossfjorden, Svalbard 79.06°N 47 Lithothamnion glaciale 169.8 this study Krossfjorden, Svalbard 79.10°N 50 Lithothamnion glaciale 181.5 this study Krossfjorden, Svalbard 79.10°N 41 Lithothamnion glaciale 181.5 this study Mosselbukta, Svalbard 79.90°N 44 Lithothamnion glaciale 119.8 this study Nordkappbukta, Svalbard 80.53°N 27 Lithothamnion glaciale 157.7 this study Nordkappbukta, Svalbard 80.53°N 45 Lithothamnion glaciale 105.3 this study Nordkappbukta, Svalbard 80.53°N 38 Lithothamnion glaciale 100.9 this study Gulf of Chiriquí, Panama 07.60°N 12–53 Lithothamnion sp. 81.09 Schäfer et al. (2011) Gulf of Panama, Panama 08.00°N 3–26 Lithothamnion sp. 23.38 Schäfer et al. (2011) Mannin Bay, Ireland 53.46°N <10 Lithothamnion corallioides 29–164 Bosence (1980) Mannin Bay, Ireland 53.46°N <10 Phymatolithon calcareum 79–249 Bosence (1980) Mannin Bay, Ireland 53.46°N <10 Lithothamnion corallioides 212–1197 Bosence & Wilson (2003) Bay of Brest, France 48.35°N 0–10 Lithothamnion corallioides 876 Potin et al. (1990) Bay of Brest, France 48.33°N 1–10 Lithothamnion corallioides 150–3000 Martin et al. (2006) Storvoll Reef, Norway 70.00°N 7 Lithothamnion cf. glaciale 895–1432 Freiwald & Henrich (1994) Straumen Bioherm, Norway 69.67°N 18–20 Lithothamnion cf. glaciale 420–630 Freiwald & Henrich (1994) Manorbier, GB 51.64°N <1 Lithophyllum incrustans 378.96 Edyvean & Ford (1987) West Angle Bay, GB 51.69°N <1 Lithophyllum incrustans 59.76 Edyvean & Ford (1987) Lizard Island, Australia 14.67°S 0 Hydrolithon onkodes 10,300 Chisholm (2000) Lizard Island, Australia 14.67°S 18 Neogoniolithon conicum 1500 Chisholm (2000) Arvoredo Island, Brazil 27.25°S 7–20 Lithophyllum sp. 55.0–136.3 Gherardi (2004)

91 Discussion

Since rhodoliths consisting of L. glaciale constitute the majority at all four sites and because P. tenue does not produce protuberances (Düwel & Wegeberg 1996), specimens consisting of P. tenue were excluded from the measurements. This ensured that the initial parameters for the calculations from each site were the same, resulting in a certain number of examined rhodoliths per station (Table 7.4). The protuberances on the upper, living side of these rhodoliths were counted and, together with the measured rhodolith surface, resulted in the median number of protuberances per square metre (Table 7.5). These numbers of protuberances per square metre do not show a particular pattern related to the water depth or the geographical latitude, which contrasts the findings of Freiwald (1993), where the number of protuberances considerably rises with increasing water depth. This is presumably owed to the large distances between the particular sampling sites at Svalbard, implying that other influencing parameters superimpose the water depth pattern. Nevertheless, Levene’s test for homogeneity of variance indicates a high significance for the coherence between the rhodolith surface and the particular number of protuberances. The coralline algae are the most consistently and heavily calcified group of the red algae and as such have been elevated to ordinal status (Corallinales Silva & Johansen, 1986). Their calcification process involves high magnesium calcite precipitation within most cell walls (Bosence 1991). According to Alexandersson (1974; 1977) and Okazaki et al. (1982), coralline algae induce a microenvironment suitable for carbonate precipitation by metabolic excretion of alginic acid, so that the calcite saturation of the ambient water column is of minor importance for the calcification process. Early electron microscope work showed that the calcified cell walls of coralline algae have a two-layered structure, an inner layer of acicular calcite parallel to the cell wall, succeeded by radial, inward growing calcite crystals (Bailey & Bisalpu- tra 1970; Alexandersson 1974; 1977; Flajs 1977a; b; Garbary 1978; Cabioch & Giraud 1986). Microprobe plotting of magnesium and calcium concentrations by Flajs (1977b) and Massieux et al. (1983) indicated that the secondary radial layer has a higher magnesium content than the earlier parallel layer. These findings are in line with the EDX-based Mg/Ca ratio measurements from this study, where a longitudinal section through a protuberance of L. glaciale shows a distinct pattern of growth increments, at which each increment consists of a varying number of cell rows (Figs 7.27; 7.28). The increments start with dense, heavily calcified cell rows, grading into less calcified cell rows. Since biomineralisation proceeds throughout the growth period, the earlier formed cell rows are further calcified than cell rows formed towards the end of the growth period. The ongoing calcification coincides with the production of the secondary cell layer (Bosence 1991), which has a higher magnesium content than the earlier parallel layer (Flajs 1977b; Massieux et al. 1983), so the Mg/Ca ratio results in a correlation of high Mg/Ca values with heavily calcified cell walls and low Mg/Ca values with less calcified cell walls (Fig. 7.27). This gives rise to an annual banding pattern exposed in the protuberances of L. glaciale. There have been a lot of discussions on the nature of the coralline algal banding patterns, ranging from daily over lunar to annual cycles. Agegian (1981) reports on growth bands of primary and secondary order in specimens of Porolithon gardineri (Foslie) Foslie, 1909 [unaccepted; accepted as Hydrolithon gardineri (Foslie) Verheij & Prud’homme van Reine,

92 Discussion

1993]. Here, the primary growth bands consisitng of one cell row each have been shown experimentally to be daily. The secondary growth bands are presumed to occur monthly, following a lunar cycle, and differ in their skeletal density. Bosence (1980) describes annual growth increments for Lithothamnion corallioides (Cr- ouan & Crouan) Crouan & Crouan, 1867 and Phymatolithon calcareum (Pallas) Adey & McKibbin, 1970. The increments are presumed to result from slow winter growth with few cell divisions and faster summer growth with numerous cell divisions, leading to elongated winter and short summer cells. Freiwald (1993) distinguishes three orders of growth increments for specimens of Lithothamnion glaciale. While the first order increments are represented by one cell row, the second order increments are presumed to be lunar or rather tidal. The algae derive from northern Norway (c. 70°N) and are hence subject to a distinct seasonality of the light regime, resulting in third order increments that consist of one dark and one bright band (see also Fig. 4.3). Freiwald & Henrich (1994) point out that this pattern represents annual increments comprising one dark band formed during winter and one bright band formed during summer. The colour of the dark bands results from a increased production of glycoproteins that coincides with an increased nutrient amount in the water column during the winter time (Freiwald 1995). These areas of intensified glycoprotein production can be stained with fuchsine solution, that further amplifies the visible banding pattern (Frei- wald 1995). The influence of the distinct seasonality of the light regime also holds for the rhodoliths from this study. Because the growth increments show a similar pattern of dark and bright bandings (Fig. 7.28) as in the study of Freiwald (1993) and also belong to same species (Lithothamnion glaciale), and according to the findings discussed above, the growth increments of the Sval- bard rhodoliths can be considerated to be annual. An additional, very striking feature is the scatter of the reproductive conceptacles, which contain the spores and form annually (Jackson 2003). These conceptacles coincide with the growth increments, meaning that one row of conceptacles is formed per growth increment (Fig. 7.28), strengthening the assumption of an annual banding pattern and giving the opportunity to estimate the annual growth of one protuberance. Since the reduced major axis algorithm and the Levene’s test for homogeneity also indicate that the annual weight increase is significantly linear (Fig. 7.29), protuberances of different age can be used for the calculation of the annual carbonate production by the rhodoliths. After the fuchsine-staining was conducted, the increments could be counted easily with the help of the annual forming reproductive conceptacles. This method led to the annual carbonate production of the rhodoliths per square metre per year (Table 7.7). The values significantly differ from each other with respect to the different sites, giving rise to the assumption that the carbonate production by the rhodolith beds is strongly influenced by physical parameters, since the regarding coralline algae belong to the same species. The other consistent characteristic for all sites is the water depth the samples derive from, except for sation 714 in 27 m water depth, that exhibits relatively high carbonate production rates, presumably due to the increased irradiance levels in those shallower water depths. This is also

93 Discussion the reason why site number 714 was excluded from the considerations regarding the possibly influencing parameters. Within these considerations, the following parameters will be handled: Geographical latitude, mean duration of sea ice cover, duration of the polar night, annual mean temperature, water depth, and calcite saturation of the water column (Table 7.8, Fig. 7.30). The first three mentioned parameters, geographical latitude, mean duration of sea ice cover, and duration of the polar night, show very clear coherences using reduced major axis algorithm and Levene’s test for homogeneity indicates high significances (Fig. 7.30a-c). This is not astonishing, because all three parameters are directly linked with the most important feature for the coralline algae, the irradiance (Kain & Norton 1990). Due to the decline of the Earth’s axis, the duration of the polar night prolongs wih increasing geographical latitude, leading to a shortage of the growth period towards higher latitudes. This is directly reflected in the amount of carbonate production, because the biomineralization process is presumed to take place only during the growth period (Bosence 1991), which also results in the banding pattern, starting with heavily calcified cell rows at the beginning of the growth period and grading into less calcified cell rows towards the end of the growth period. The same applies for the mean duration of the sea ice cover, since ice cover reduces the PAR transmitted through the water column to a minimum and if the sea ice is covered with a snow layer, the increased albedo reduces the PAR reaching the coralline algae to a minimum. Despite the high significance between the carbonate production of the rhodoliths and the mean duration of the sea ice cover, the correlation has to be considered conservative, because the AMSR-E Sea Ice data base only provides information for the last ten years, while the oldest examined rhodoliths expose maximum ages of up to >90 years (Fig. 7.29; station 701). Even considering that, the characteristics clearly indicate the strong coherence between the carbonate production by the rhodoliths and geographical parameters. On the other hand, it has to be mentioned that also the polar day prolongs with increasing geographical latitude, so the question is why both effects do not overturn against each other. The reason is that the photosynthesis of the coralline algae already starts at very low PAR- levels (Ragan 1981; Roberts et al. 2002), i.e. the carbon production period of the algae starts much earlier than that of many other plants, giving them a head start also in the accumulation of starch as a carbon reservoir (Fig. 7.20). This is also possible because of the macro-nutrients like nitrogen and phosphorus, which increasingly deplete during the summer season but are storaged by the coralline algae during their availability in the winter season (see above). With increasing daylength and hence a raised PAR, the advantage of the coralline algae over many green plants successively decreases, since the irradiance levels now give a competitive edge to the green, faster growing plants. The next physical parameter showing a high coherence with the annual carbonate production of the rhodoliths is the annual mean temperature (Fig. 7.30d). As stated before, the optimum temperature clearly varies geographically and with species, but the general pattern usually shows an increase in growth rate to a maximum that is near the top of the tolerated range (Kain & Norton 1990). This is directly reflected in the carbonate calculations, where the annual carbonate production decreases linear to the annual mean temperature, showing a high

94 Discussion significance using Levene’s test for homogeneity. The annual mean temperatures decrease towards the northern sites since the influence of the warm AW transported by the WSC [whose core temperatures range from 6-8°C and the salinities from 35.1-35.3 (Aagaard et al. 1987)] decreases in favour of the colder ArW transported by the ESC [whose core temperatures range from 1-3°C and the salinities from 34.5-34.9 (OSPAR Commission 2000)], and both currents have a high impact on the local climate (Orvik & Niiler 2002; Sapota et al. 2009). While the effects of the previously mentioned parameters are high, water depth and calcite saturation of the water column do not seem to have an influence on the carbonate production by the rhodoliths (Fig. 7.30e-f). The missing influence of water depth is easy to explain, since the depths are nearly the same, all ranging in the dysphotic zone (Table 7.8; Fig. 7.13), except for station 714 at Nordkappbukta, which has been excluded from the calculations. Hence, the effect of the slightly varying water depths is superimposed by far by the effects mentioned above. One can assume that, if the samples would derive from the same site, differing only in the water depth they would have been collected, water depth would have marked effects on the annual carbonate productions due to its great influence on the PAR available to the rhodoliths. There is also no reasonable coherence between the anual carbonate production of the rhodoliths and the calcite saturation of the water column. Regarding the regression line in Fig. 7.30f, the annual carbonate production seems to increase with decreasing calcite saturation, which would contrast all assumptions with respect to the biomineralization process of the coralline algae. As Alexandersson (1974; 1977) and Okazaki et al. (1982) state, coralline algae induce a microenvironment suitable for carbonate precipitation by metabolic excretion of alginic acid, so that the carbonate saturation of the ambient water column is of minor importance for the calcification process, which should hold for as long as the water column is still at least slightly saturated with respect to calcite (ΩCal ≥1), so the algae can deploy the free calcium ions. Hence, as it is true for the water depth, the effect of the calcite saturation of the water column is strongly superimposed by the effects assiociated with the geographical latitude, the duration of the polar night, the duration of the sea ice cover, and the annual mean temperature. Though, in case of calcite undersaturation (ΩCal <1), the impact on coralline algal carbonate production rates is likely to increase, as Martin et al. (2008) showed experimentally that an increasing acidification of seawater leads to a significant reduction in coralline algal cover, indicating that a lowered pH and a significant reduction of carbonate saturation are important factors affecting the rhodolith beds.

With respect to the ongoing climate change, one can say that it will have distinctive impacts on the annual carbonate production of the rhodoliths. There will be no change in the duration of the polar night, since this is subject to astronomic parameters. But regarding the duration of the sea ice cover and the annual mean temperature, a weakening of the Gulf Stream, as it is proposed to parallel the ongoing global warming (Quadfasel 2005), would have strong impact. This would result in a decrease of the annual mean temperature, since the influence of the cold ArW transported by the ESC would prevail throughout the year, thus disadvantaging the growth processes of the coralline algae (Kain & Norton 1990). It may also delay the breakup of the annual forming sea ice, thus prolonging the dark period. Indeed it has been shown that coralline algae can cope with several months of complete darkness before respiration suddenly

95 Discussion drops (Lüder et al. 2002), but the aftermath on specimens of L. glaciale and P. tenue have not been under examination yet. The weakening of the Gulf Stream as it is proposed by Quadfa- sel (2005) is only one of many scenarios. Another and more probable development, as it can already be observed, is the increasing disappearence of sea ice due to rising temperatures. This was confirmed by Shapiro et al. (2003), showing the retreat of the mean ice edge position in the Barents Sea over a 152-year period from 1850 to present.The outcome for the rhodolith communities of such a development is not predictable, but due to the climatic changes, it may result in conditions similar to those described for the rhodolith communities in northern Norway (Freiwald 1993; Freiwald & Henrich 1994), including the invasion of various species. At first view, this does not seem harmful to the rhodolith communities, because even the rhodoliths in northern Norway, which also mainly consist of L. glaciale specimens, thrive well under the current conditions. But it has to be noticed that the production rates of the Norwegian rhodoliths are much higher due to thicker annual growth increments (annual carbonate productions of 895-1432 g m-2 yr-1 and 420-630 g m-2 yr-1; see also Table 8.1; Freiwald 1993), resulting principally from the shallower water depth (7 m and 18-20 m, respectively; Freiwald 1993) and the shorter duration of the polar night (53 days; data from USNO Sun Rise Tables). Regarding the resulting, longer duration of the growth period for the coralline algae and the higher production rates, it gets clear that the residing algae can better cope with the grazing pressure exerted by echinoids and chitons than it was the case for the Svalbard communities, if they where increasingly affected by the Norwegian grazers. With respect to the calcite saturation of the water column, another upcoming problem was the ongoing ocean acidification. Coralline algae induce a microenvironment suitable for carbonate precipitation and hence are quite independet of the conditions in the water column (Alexandersson 1974; 1977; Okazaki et al. 1982), but regarding the experiments by Martin et al. (2008; see above), this should only hold as long as the water is least slighlty saturated with respect to calcite (ΩCal ≥ 1). Additionally, if anthropogenic CO2 emissions follow the IPCC business as usual scenario (SRES A2; Steinacher et al. 2009), it has been shown that this could lead to a decrease in rhodolith abundance, because coralline algae are very sensitive to risen CO2 conditions and the coequally lowered pH, leading to a strong decrease in the percentage of area covered (Hall-Spencer et al. 2008; Martin et al. 2008). Altogether, coralline algae can thrive under a broad range of carbonate saturation values, but undersaturated conditions would have a negative impact and presumably decrease the carbonate production rates of the rhodolith beds. The rhodolith forming coralline algae L. glaciale and P. tenue are important carbonate producers at the four investigated Svalbard localities, along with barnacles and molluscs, thus representing a unique polar carbonate factory. If the climate change is going to proceed, these highly specialised organisms are assumed to be strongly affected. Since they have an important role as ecosystem engineers on the otherwise relatively shelterless glaciogenic flats, a decrease in their abundance will also have an unknown impact on the associated benthic organisms.

96 Discussion

8.4 Comparison with other rhodolith communities

Beside the sites examined in this study, other polar and subpolar rhodolith communities occur, for example, in Alaska (Konar & Iken 2005; Konar et al. 2006) and mainland Norway (Freiwald 1993; 1998; Freiwald & Henrich 1994). The Alaskan community, situated in Herring Bay (Prince William Sound) at 60°28’N and 47°45’W, is the northernmost known in the Pacific Ocean. In contrast to the Svalbard communities, Herring Bay features a monospecific rhodolith community composed of Phymato- lithon calcareum (Pallas) Adey & McKibbin, 1970. While the seafloor at the Svalbard sites consists almost completely of presumably glaciogenic pebbles, cobbles, and boulders with fine sediments restricted to patches, the seafloor at Her- ring Bay showed that 60% of the substrate was hard boulders and 40% fine sediments (Konar & Iken 2005). In both cases, the occurrence of coralline algae is restricted to the hard substrate, implying that the competition for space between the coralline algae and other fixosessile organisms is more distinctive at Herring Bay than at the Svalbard sites. Konar & Iken (2005) showed that, although the coralline algae are the major space occupiers (>60%), they are not the competitive dominants against many other sessile organisms. This may be due to the high abundance of bioeroding chitons (Konar et al. 2006), that have a stronger impact on the other sessile organisms like sponges, bryozoans and tunicates (Konar & Iken 2005) than on the coralline algae, that possess various adaptations against bioerosion (Steneck 1986). Regarding that, it becomes clear that the Herring Bay community seems to show the same interaction of coralline algae and bioeroders as in Svalbard and other localities (Steneck 1986). The observations by Konar & Iken (2005) also showed that the Alaskan rhodoliths have a similar ecological function like those in Svalbard as they provide a habitat for the benthic organisms, and their branches are inhabited by a specialised cryptofauna. This causes a strong increase in diversity of the micro- and macro-benthos, an attribute that seems to characterise many rhodolith communities (Foster 2001). In Norway, the Storvoll Plateau (Freiwald 1993; 1998; Freiwald & Henrich 1994), situated in the Troms district at the southern tip of Rebbenesøy (69°59’N; 18°40’E), consists of rhodolith communities that fringe a rigid, in situ red algal buildup in 14-15 m water depth. The community is multispecific, consisting of Lithothamnion cf. glaciale, Lithothamnion sp., and Phymatolithon sp., but the proportion of the two genera is much more balanced than in Svalbard. However, the major sources for rhodolith production are detached heads from Litho- thamnion-branches, so most of the rhodoliths are not nucleated as they are in Svalbard. Be- cause of that, the shape of the rhodoliths can be used as an indicator for the hydrodynamic regime and the findings of Freiwald (1993) show a strong concentration on spheroidal rhodolith shapes (88%) at the Straumensund, indicating high energy hydrodynamics (Bosence 1983a; Freiwald 1993). Contrariwise, the shapes of the rhodoliths from Svalbard cannot be used as indicators for the prevailing hydrodynamic regime. Since all rhodoliths are nucleated, their shape is widely predetermined by their particular nucleus (Freiwald 1995). The lithoclastic nuclei, that are

97 Discussion encrusted by coralline algae, however, may be used as indicators for former hydrodynamic conditions (Sneed & Folk 1958; Bloore 1977; see above). Moreover, the Storvoll Plateau community (14-15 m water depth) occurs in considerably shallower water than the Svalbardian communities (mostly 30-75 m water depth). This, beside the shorter duration of the polar night (53 days; data from USNO Sun Rise Tables) may also be the main reason for the higher carbonate production rates at Norway compared to the Svalbard sites. Though the rhodolith assemblage is composed of L. glaciale to a significant percentage, the mean annual carbonate productions amount to 841 g m-2 yr-1 in 7 m water depth at the Storvoll Plateau (Freiwald 1993) compared to 200.3 g m-2 yr-1 in 45 m water depth at Floskjeret, being the Svalbard site with the highest carbonate production rate. The benthic organisms inhabiting the Norwegian rhodolith communities are similarly composed compared to the Svalbard sites, and many species (e.g. Chlamys islandica, Hiatella arctica, and Hemithiris psittacea) occur at both localities. Even bioeroders like Strongylocentrotus droebachiensis (Müller, 1776) and Lepidopleurus asellus Gmelin, 1791 are common and take the same ecological niche as the grazers in Herring Bay and Svalbard do. This further confirms the assumptions of Steneck (1986), that coralline algae require the presence of bioeroding organisms to keep themselves clean and healthy, since otherwise they would be overgrown by competitive fixosessile invertebrates (Steneck 1986; Konar & Iken 2005), which would lead to the successive dieback of the coralline algae.

Overall, these rhodolith assemblages are ecologically very important, especially in providing substratum and protection for many benthic organisms, but also food resources for the grazers. This system works similarly at the different sites, even if the communities are composed of different species, which however fulfil the same functions. If rhodoliths are present in bio- cenoses like the Svalbard sites, which mainly consist of flat gravel pavements, they can greatly increase the diversity by providing a kind of microenvironment for cryptofauna between their branches (Steller et al. 2003; Konar & Iken 2005) and in their partly hollow bodies. Co- evally, several species present between and on the rhodoliths (e.g. chitons and echinoids) act as grazers and keep the corallines free from epiphytes (Steneck 1986).

98 Discussion

8.5 Future implications

The Svalbard rhodolith communities are highly specialised in their adaptations to the physical environment as well as in their interaction with the present fauna, showing parallels to other polar rhodolith communities (Freiwald 1993; Freiwald & Henrich 1994; Konar & Iken 2005; Konar et al. 2006). Climatic change will likely result in shifts in light conditions, temperatures and sedimentation processes or the development of carbonate undersaturation caused by ocean acidification. Therefore, it will also have distinct impacts on the rhodolith communities. The coralline algae prevailing at the four sites depend on a small window in the temperature range. If the winter mean temperatures would decrease, the vital processes may be disrupted (Kain & Norton 1990), which would lead to a deferred start at the beginning of the growing season, or even to a complete dieback of the thalli. On the other hand, strongly increasing mean temperatures would presumably affect the formation of reproductive conceptacles during the growing season (Hall-Spencer 1994), so the settling gametes were not able to leverage the enhanced conditions of the light regime and hence would go under other, faster growing plants. Fluctuations in salinity may cause osmotic stress, unfavourable ionic balances and a shortage of essential metabolites (Kain & Norton 1990). Since the main coralline algal carbonate producer, L. glaciale, is also able to tolerate both low (<18, Wilson et al. 2004) and strongly fluctuating salinities (18-40 in sea lochs off the west coast of Scotland, Jackson 2003), the seasonal decreasing salinities induced by melting glaciers during summer time likely have non or at most low direct impact on the rhodolith communities. Though, it should be stressed that L. glaciale indeed tolerates low and fluctuating salinities but is best adapted to fully marine conditions (Wilson et al. 2004). This implies that permanently decreased salinities also in deeper water layers, as it may be a consequence of changing hydrography (Svendsen et al. 2002; Clarke & Harris 2003), would negatively affect the rhodolith beds and lead to impaired growth conditions. Irradiance is another factor that limits the appearance of coralline algae at Svalbard to a distinct depth gradient, since light conditions seem to be insufficient in water depths >78 m, while in a water depth <30 m, irradiance is high enough to favour other plants, which displace the coralline algae. Regarding that, the outcome of a potential global warming for the light regime and hence the thriving of the coralline algae at Svalbard is ambiguous. On the one hand, increasing temperatures would lead to shorter periods of sea ice cover and hence prolong the growing season of the coralline algae. On the other hand, this would also affect faster growing plants without low-light-adaption, leading to an increased competition for space. A major problem arising with increasing annual mean temperatures was the simultaneous glacier meltdown, resulting in the transport of huge amounts of fine sediments to the water column (Konar & Iken 2005). This would strongly impair the growth conditions for the coralline algae due to a reduced PAR and the parallel burial with fine sediment, being a major reason for the dieback of coralline algae (Wilson et al. 2004).

99 Discussion

The concomitant appearance of coralline algae and prominent grazers, such as Tonicella rubra and Strongylocentrotus sp., keeps the coralline algae free from epiphytes and coequally provides feeding grounds for the grazers by means of the biofilms present on the surface of the rhodoliths (Steneck 1986). Additionally, the rhodolith accumulations act as bioengineers and represent microenvironments on the otherwise non-protected glaciogenic flats, with hollow rhodoliths being of particular significance as providers of microhabitats for the associated benthic fauna. A regression in the rhodolith stock due to a dieback of the coralline algae would presumably result in more unfavourable conditions for many benthic organisms due to the loss of habitat and food resources. The importance of at least slight carbonate saturation for coralline algae is well known, and observations by Hall-Spencer et al. (2008) showed that decreased pH levels can result in the rapid decline of calcareous algae. Additionally, the abundances of gastropods and sea urchins, whose function as surface grazers is essential for the thriving of coralline algae (Steneck 1985; 1986), may be negatively affected by lower pH levels (Hall-Spencer et al. 2008). The current state of the seawater carbonate system is suitable for the rhodolith communities. However, high-latitude oceans will very likely be the first to become undersaturated with respect to calcite and aragonite (Orr et al. 2005), and modelling studies project annual mean carbonate undersaturation as soon as 2032 (Steinacher et al. 2009). This development will have a massive impact on the rhodoliths and many associated benthic organisms. On the one hand, a decline in rhodolith coverage can lead to impaired feeding and breeding conditions for the associated grazers, and on the other hand, reduced abundances of grazing organisms can result in an excessive overgrowth of rhodoliths by epiphytic algae (Steneck 1986). Overall, Lithothamnion glaciale and Phymatolithon tenue appear to be well adapted to the extreme environment of the Arctic. But like all ecosystems with highly specialised organisms present, even this environment is vulnerable to global change. The findings reveal that coralline algae are much more widespread in polar waters than previously thought, thus representing a polar carbonate factory of significant importance. In contrast to other polar carbonate facories, which feature heterotrophic filter-feeders as main carbonate producers (Henrich et al. 1992; Henrich et al. 1995; Andruleit et al. 1996;), the Svalbard communities are strongly dominated by photoautotrophic frame builders, thus being particularly sensitive to future changes in the light regime.

100 Conclusions

9. Conclusions

The findings of this study revealed various aspects of the rhodolith communities at the Sval- bard archipelago, concerning their physical and biological controls, their significance as polar biosedimentary system, and their potential susceptibility to environmental changes. These findings are summarised as follows.

• The shapes of the rhodoliths cannot be used as indicators for the prevailing hydrodynamic regime since all rhodoliths are nucleated or presumably contained a nucleus at an earlier growth stage. However, the shape of the nuclei may be used as an indicator for former hydrodynamics and the onset of conditions favourable to rhodoliths may be approximately determined by the age of the rhodolith communities.

• The presence of an appropriate substrate is essential for the thriving of coralline algae. The substrate has to offer a minimum stability and must not be smothered by fine sediments. It is one of the main controls susceptible to environmental changes.

• Regarding the Svalbard archipelago, present-day temperatures and salinities are suitable to the dominating coralline algal species, Lithothamnion glaciale. Significant changes, especially increasing temperatures and decreasing salinities, would presumably lead to impaired conditions for the rhodolith communities.

• The irradiance is the most important control on the coralline algae and limits their appearance to a distinct depth gradient, principally situated in the dysphotic zone.

• The coralline algae can cope with shifts of the carbonate saturation in the water column, as long as the water is at least slightly saturated with respect to calcite. How- ever, increasing ocean acidification will have a strong impact on the rhodolith communities.

• The rhodoliths are an important part of the prevailing ecosystem. They act as ecosystem engineers, and the biofilms on their surface represent a major source of food for many grazing organisms. Coequally, the rhodoliths benefit from the cleaning effect exerted by the grazers.

• The rhodolith beds are an important polar carbonate factory, along with barnacles and molluscs. Being photosynthetic organisms, their carbonate production rates significantly decrease towards higher latitudes, mainly as a response to the prolonged polar night.

101 References

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116 Online data sources

Online data sources

Svalbards present day population

Data base Statistics Norway; comprising all official statistics in Norway.

Available from: http://www.ssb.no/english/

Mean water temperatures and salinities

Data base LEVITUS 94; comprising the World Ocean Atlas 1994 (WOA94), an atlas of objectively analyzed fields of major ocean parameters at the annual, seasonal, and monthly time scales.

Available from: http://iridl.ldeo.columbia.edu/SOURCES/.LEVITUS94/

Annual sea ice formation and breakup

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Available from: http://www.iup.uni-bremen.de:8084/amsr/

Duration of the Polar night

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117 Appendix A

Appendix A

Raw data CTD-measurements [Note: Data collected by A. Form ([email protected]) and M. Meyerhöfer ([email protected]) during MSM 02/03 expedition of RV Maria S. Merian in 2006]

Station 616 Station 632 Station 640 Station 641 Station 642

Depth Temperature Salinity Depth Temperature Salinity Depth Temperature Salinity Depth Temperature Salinity Depth Temperature Salinity [m] [°C] (PSS) [m] [°C] (PSS) [m] [°C] (PSS) [m] [°C] (PSS) [m] [°C] (PSS)

2 5.784 32.022 2 6.321 32.576 2 3.564 31.229 3 5.193 32.589 3 5.703 31.051

3 5.764 32.025 3 6.286 32.605 3 3.601 31.940 4 5.194 32.710 4 5.746 31.585

4 5.744 32.021 4 6.246 32.770 4 3.611 31.95 5 5.199 32.730 5 5.778 31.660

5 5.732 32.020 5 6.276 32.652 5 3.635 32.032 6 5.213 32.756 6 5.785 31.730

6 5.735 32.020 6 6.298 32.625 6 3.643 32.076 7 5.099 32.813 7 5.786 31.740

7 5.726 32.019 7 6.253 32.767 7 3.680 32.211 8 5.209 32.840 8 5.787 31.768

8 5.715 32.018 8 6.233 32.828 8 3.732 32.412 9 5.212 32.855 9 5.793 31.847

9 5.673 32.016 9 6.209 32.914 9 3.803 32.651 10 5.200 32.852 10 6.049 32.735

10 5.667 32.015 10 6.159 33.144 10 3.825 32.743 11 5.040 32.849 11 6.304 33.024

11 5.537 32.058 11 6.162 33.182 11 3.896 32.890 12 5.252 32.987 12 6.395 33.123

12 4.779 32.447 12 6.164 33.230 12 4.123 33.179 13 5.276 33.076 13 6.081 33.133

13 4.401 32.579 13 6.159 33.269 13 4.121 33.215 14 3.701 33.053 14 6.115 33.178

14 4.162 32.667 14 6.115 33.340 14 3.889 33.221 15 3.715 33.120 15 6.158 33.205

15 3.850 32.770 15 6.064 33.399 15 3.944 33.227 16 4.438 33.315 16 6.097 33.204

16 3.621 32.847 16 6.026 33.447 16 3.907 33.269 17 4.907 33.460 17 6.001 33.254

17 3.542 32.942 17 5.979 33.506 17 3.968 33.313 18 4.979 33.501 18 6.146 33.361

18 3.411 33.031 18 5.978 33.507 18 4.075 33.330 19 4.648 33.580 19 5.893 33.686

19 3.366 33.154 19 5.961 33.557 19 4.656 33.489 20 4.436 33.652 20 5.797 33.798

20 3.401 33.223 20 5.933 33.626 20 4.958 33.570 21 4.246 33.704 21 5.845 33.858

21 3.395 33.221 21 5.916 33.687 21 5.110 33.610 22 4.303 33.759 22 5.813 33.874

22 3.503 33.340 22 5.893 33.779 22 5.362 33.676 23 4.182 33.783 23 5.796 33.886

23 3.573 33.422 23 5.839 33.858 23 5.322 33.688 24 4.507 33.866 24 5.709 33.901

24 3.614 33.454 24 5.815 33.884 24 5.196 33.688 25 4.374 33.868 25 5.614 33.919

25 3.682 33.565 25 5.786 33.909 25 5.087 33.681 26 4.494 33.921 26 5.580 33.941

26 3.692 33.593 26 5.773 33.924 26 5.048 33.675 27 4.391 33.935 27 5.480 33.992

27 3.667 33.630 27 5.721 33.986 27 4.063 33.603 28 4.530 33.977 28 5.461 34.024

28 3.616 33.685 28 5.730 34.105 28 3.054 33.591 29 4.659 34.016 29 5.408 34.030

29 3.623 33.774 29 5.660 34.143 29 3.312 33.743 30 4.681 34.045 30 5.078 34.042

30 3.613 33.818 30 5.614 34.185 30 3.748 33.949 31 4.538 34.074 31 5.170 34.075

31 3.539 33.837 31 5.544 34.202 31 3.936 34.037 32 4.664 34.121 32 4.829 34.091

32 3.429 33.847 32 5.512 34.240 32 4.026 34.071 33 4.757 34.159 33 4.523 34.120

33 3.442 33.899 33 5.525 34.265 33 4.144 34.125 34 4.688 34.161 34 4.155 34.141

34 3.650 33.971 34 5.541 34.284 34 4.167 34.191 35 4.630 34.188 35 4.050 34.184

35 3.559 34.000 35 5.552 34.337 35 4.172 34.262 36 4.550 34.219 36 4.224 34.243

36 3.623 34.028 36 5.548 34.371 36 3.880 34.258 37 4.311 34.219 37 4.035 34.241

37 3.628 34.06 37 5.577 34.391 37 3.676 34.241 38 3.976 34.217 38 3.846 34.249

38 3.589 34.091 38 5.585 34.427 38 3.643 34.244 39 4.046 34.256 39 3.570 34.264

39 3.603 34.104 39 5.547 34.430 39 3.399 34.234 40 4.112 34.277 40 3.527 34.293

40 3.651 34.113 40 5.554 34.447 40 3.055 34.220 41 4.131 34.292 41 2.734 34.258

41 3.652 34.124 41 5.560 34.476 41 2.766 34.205 42 3.959 34.293 42 2.553 34.279

42 3.606 34.138 42 5.530 34.519 42 2.299 34.214 43 3.740 34.286 43 2.695 34.329

43 3.577 34.150 43 5.515 34.571 43 2.330 34.229 44 3.356 34.279 44 2.770 34.357

44 3.520 34.181 44 5.483 34.613 44 2.230 34.233 45 3.25 34.299 45 2.567 34.361

45 3.500 34.189 45 5.425 34.638 45 2.250 34.241 46 3.219 34.304 46 2.406 34.360

46 3.473 34.198 46 5.399 34.653 46 2.400 34.277 47 3.162 34.312 47 2.435 34.369

47 3.469 34.199 47 5.145 34.655 47 2.680 34.336 48 3.119 34.334 48 2.344 34.373

48 3.454 34.204 48 5.238 34.681 48 2.718 34.349 49 2.874 34.341 49 2.372 34.386 Appendix A

Station 616 Station 632 Station 640 Station 641 Station 642

49 3.400 34.225 49 5.249 34.692 49 2.663 34.349 50 2.849 34.376 50 2.401 34.399

50 3.383 34.253 50 5.236 34.693 50 2.631 34.369 51 2.940 34.409 51 2.375 34.406

51 3.384 34.279 51 5.204 34.690 51 2.652 34.386 52 2.672 34.409 52 2.396 34.413

52 3.345 34.301 52 5.094 34.682 52 2.541 34.386 53 2.587 34.424 53 2.366 34.414

53 3.302 34.304 53 5.03 34.687 53 2.508 34.385 54 2.634 34.434 54 2.370 34.418

54 3.226 34.308 54 5.044 34.692 54 2.600 34.399 55 2.649 34.448 55 2.366 34.419

55 3.209 34.309 55 5.043 34.706 55 2.644 34.410 56 2.626 34.457 56 2.354 34.421

56 3.164 34.306 56 4.943 34.720 56 2.522 34.407 57 2.545 34.459 57 2.336 34.425

57 3.142 34.308 57 4.882 34.724 57 2.221 34.400 58 2.514 34.461 58 2.374 34.438

58 3.094 34.306 58 4.856 34.727 58 2.020 34.392 59 2.503 34.464 59 2.389 34.451

59 2.975 34.319 59 4.719 34.733 59 1.948 34.397 60 2.515 34.469 60 2.356 34.451

60 2.964 34.323 60 4.716 34.736 60 2.014 34.413 61 2.441 34.468 61 2.330 34.450

61 2.939 34.329 61 4.755 34.745 61 1.999 34.413 62 2.190 34.458 62 2.505 34.487

62 2.912 34.332 62 4.803 34.758 62 1.960 34.415 63 2.040 34.455 63 2.523 34.492

63 2.810 34.346 63 4.883 34.783 63 1.937 34.423 64 2.006 34.459 64 2.561 34.498

64 2.713 34.374 64 4.946 34.812 64 1.864 34.431 65 1.966 34.460 65 2.547 34.504

65 2.634 34.382 65 4.963 34.825 65 1.861 34.437 66 1.863 34.459 66 2.544 34.513

66 2.566 34.385 66 4.960 34.829 66 1.818 34.438 67 1.860 34.464 67 2.479 34.511

67 2.486 34.394 67 4.952 34.832 67 1.677 34.435 68 1.934 34.476 68 2.249 34.507

68 2.568 34.414 68 4.944 34.834 68 1.608 34.431 69 1.968 34.486 69 2.191 34.503

69 2.758 34.454 69 4.936 34.837 69 1.605 34.432 70 1.789 34.485 70 2.078 34.495

70 2.97 34.473 70 4.924 34.841 70 1.578 34.431 71 1.734 34.488 71 1.991 34.489

71 2.951 34.478 71 4.900 34.848 71 1.563 34.431 72 1.734 34.488 72 2.031 34.493

72 3.173 34.529 72 4.863 34.856 72 1.532 34.431 73 1.736 34.488 73 2.223 34.521

73 3.596 34.575 73 4.833 34.865 73 1.543 34.432 74 1.744 34.489 74 2.162 34.528

74 3.701 34.578 74 4.775 34.876 74 1.657 34.446 75 1.741 34.488 75 1.996 34.520

75 3.698 34.584 75 4.755 34.881 75 1.613 34.449 76 1.735 34.487 76 2.191 34.543

76 3.711 34.585 76 4.748 34.888 76 1.558 34.451 77 1.725 34.487 77 2.434 34.569

77 3.790 34.610 77 4.879 34.915 77 1.610 34.456 78 1.715 34.487 78 2.626 34.590

78 3.721 34.589 78 4.924 34.924 78 1.639 34.459 79 1.698 34.488 79 2.625 34.593

79 3.391 34.583 79 4.938 34.929 79 1.674 34.465 80 1.702 34.489 80 2.784 34.614

80 3.325 34.591 80 4.955 34.936 80 1.698 34.470 81 1.722 34.493 81 2.848 34.628

81 3.459 34.618 81 4.978 34.945 81 1.691 34.475 82 1.731 34.504 82 2.747 34.626

82 3.485 34.617 82 4.993 34.950 82 1.562 34.474 83 1.724 34.508 83 2.748 34.631

83 3.477 34.619 83 5.000 34.953 83 1.492 34.472 84 1.697 34.508 84 2.782 34.638

84 3.476 34.621 84 4.983 34.952 84 1.471 34.475 85 1.694 34.508 85 2.822 34.646

85 3.503 34.637 85 4.972 34.952 85 1.456 34.476 86 1.671 34.509 86 2.839 34.655

86 3.479 34.635 86 4.971 34.953 86 1.447 34.479 87 1.648 34.510 87 2.885 34.665

87 3.459 34.657 87 4.967 34.953 87 1.431 34.480 88 1.670 34.512 88 2.894 34.668

88 3.585 34.684 88 4.954 34.952 88 1.403 34.484 89 1.677 34.512 89 3.151 34.699

89 3.586 34.685 89 4.934 34.950 89 1.399 34.484 90 1.685 34.513 90 3.221 34.706

90 3.579 34.687 90 4.932 34.950 90 1.397 34.484 91 1.690 34.520 91 3.152 34.702

91 3.581 34.691 91 4.934 34.950 91 1.396 34.485 92 1.669 34.526 92 3.110 34.699

92 3.587 34.698 92 4.934 34.951 92 1.406 34.489 93 1.691 34.529 93 3.058 34.695

93 3.622 34.708 93 4.938 34.952 93 1.458 34.500 94 1.694 34.529 94 3.354 34.736

94 3.689 34.713 94 4.952 34.954 94 1.456 34.507 95 1.705 34.531 95 3.820 34.799

95 3.828 34.743 95 4.970 34.957 95 1.430 34.509 96 1.713 34.532 96 3.816 34.805

96 3.916 34.746 96 4.993 34.960 96 1.379 34.508 97 1.724 34.535 97 3.820 34.808

97 3.818 34.736 97 5.033 34.968 97 1.376 34.510 98 1.736 34.538 98 3.818 34.809

98 3.746 34.732 98 5.131 34.987 98 1.394 34.514 99 1.746 34.542 99 3.816 34.810

99 3.645 34.713 99 5.154 34.995 99 1.402 34.520 100 1.756 34.545 100 3.813 34.811

100 3.516 34.710 100 5.151 34.997 100 1.407 34.523 101 1.850 34.556 101 3.732 34.807

101 3.531 34.714 101 5.133 34.996 101 1.408 34.524 102 1.884 34.561 102 3.684 34.804

102 3.482 34.710 102 5.143 35.000 102 1.407 34.527 103 1.889 34.563 103 3.656 34.802

103 2.563 34.593 103 5.196 35.012 103 1.403 34.532 104 1.876 34.565 104 3.605 34.798

104 2.314 34.612 104 5.218 35.021 104 1.399 34.537 105 1.840 34.565 105 3.554 34.794 Appendix A

Station 616 Station 632 Station 640 Station 641 Station 642

105 2.277 34.604 105 5.171 35.019 105 1.399 34.538 106 1.826 34.564 106 3.559 34.793

106 2.182 34.590 106 5.125 35.015 106 1.398 34.542 107 1.915 34.577 107 3.697 34.814

107 2.021 34.575 107 5.084 35.012 107 1.396 34.546 108 2.065 34.597 108 3.742 34.821

108 1.963 34.572 108 5.091 35.014 108 1.397 34.555 109 2.049 34.601 109 3.752 34.824

109 1.910 34.561 109 5.100 35.015 109 1.396 34.557 110 2.038 34.602 110 3.751 34.824

110 1.930 34.586 110 5.184 35.028 110 1.391 34.559 111 1.907 34.599 111 3.675 34.819

111 2.191 34.597 111 5.215 35.034 111 1.390 34.560 112 1.901 34.599 112 3.529 34.809

112 2.004 34.580 112 5.268 35.042 112 1.397 34.561 113 1.901 34.601 113 3.498 34.805

113 2.009 34.589 113 5.254 35.041 113 1.404 34.544 114 1.907 34.605 114 3.503 34.805

114 2.041 34.605 114 5.228 35.039 114 1.403 34.553 115 1.892 34.609 115 3.500 34.804

115 2.434 34.633 115 5.247 35.043 115 1.403 34.559 116 1.923 34.613 116 3.461 34.799

116 2.422 34.631 116 5.280 35.048 116 1.402 34.558 117 2.009 34.622 117 3.346 34.791

117 2.521 34.647 117 5.280 35.049 117 1.400 34.554 118 2.074 34.631 118 3.318 34.788

118 2.338 34.622 118 5.238 35.044 118 1.383 34.578 119 2.134 34.638 119 3.255 34.783

119 2.133 34.623 119 5.076 35.024 119 1.380 34.577 120 2.01 34.634 120 3.169 34.776

120 2.129 34.621 120 4.960 35.010 120 1.373 34.578 121 1.944 34.631 121 3.164 34.774

121 2.129 34.621 121 4.965 35.010 121 1.354 34.582 122 1.904 34.630 122 3.253 34.786

122 2.244 34.642 122 4.988 35.013 122 1.352 34.585 123 1.891 34.633 123 3.336 34.798

123 2.408 34.659 123 4.974 35.013 123 1.359 34.585 124 2.089 34.649 124 3.302 34.797

124 2.603 34.681 124 5.009 35.019 124 1.363 34.587 125 2.014 34.648 125 3.222 34.792

125 2.806 34.698 125 5.144 35.039 125 1.351 34.591 126 1.913 34.642 126 3.150 34.785

126 3.182 34.758 126 5.160 35.043 126 1.331 34.593 127 1.813 34.638 127 2.857 34.765

127 3.412 34.780 127 5.045 35.033 127 1.339 34.591 128 1.773 34.634 128 2.649 34.75

128 3.354 34.772 128 5.090 35.039 129 1.713 34.632 129 2.640 34.746

129 3.306 34.772 129 5.109 35.042 130 1.727 34.636 130 2.627 34.743

130 3.298 34.771 130 5.144 35.047 131 1.840 34.648 131 2.561 34.737

131 3.233 34.773 131 5.139 35.050 132 1.855 34.651 132 2.529 34.734

132 3.406 34.795 132 5.155 35.054 133 1.825 34.649 133 2.490 34.730

133 3.498 34.810 133 5.203 35.063 134 1.749 34.644 134 2.428 34.724

134 3.630 34.827 134 5.220 35.066 135 1.599 34.635 135 2.333 34.716

135 3.888 34.865 135 5.222 35.068 136 1.573 34.632 136 2.308 34.713

136 3.985 34.878 136 5.249 35.072 137 1.540 34.632 137 2.294 34.712

137 4.009 34.881 137 5.261 35.075 138 1.521 34.630 138 2.252 34.709

138 3.967 34.876 138 5.316 35.085 139 1.517 34.630 139 2.257 34.709

139 3.836 34.866 139 5.354 35.092 140 1.513 34.630 140 2.261 34.708

140 3.745 34.865 140 5.338 35.091 141 1.510 34.630 141 2.305 34.713

141 3.725 34.864 141 5.277 35.086 142 1.509 34.630 142 2.520 34.736

142 3.722 34.864 142 5.219 35.079 143 1.508 34.631 143 2.539 34.740

143 3.718 34.862 143 5.185 35.075 144 1.515 34.632 144 2.538 34.741

144 3.665 34.860 144 5.182 35.074 145 1.537 34.634 145 2.507 34.739

145 3.722 34.874 145 5.131 35.069 146 1.608 34.641 146 2.491 34.738

146 3.800 34.888 146 5.111 35.067 147 1.7 34.650 147 2.507 34.739

147 3.887 34.895 147 5.079 35.063 148 1.771 34.659 148 2.494 34.737

148 3.792 34.869 148 5.034 35.058 149 1.789 34.663 149 2.327 34.729

149 3.498 34.861 149 4.984 35.053 150 1.832 34.670 150 2.320 34.729

150 3.490 34.856 150 4.964 35.050 151 1.834 34.671 151 2.309 34.728

151 3.512 34.861 151 4.956 35.049 152 1.820 34.672 152 2.276 34.727

152 3.582 34.871 152 4.872 35.040 153 1.805 34.671 153 2.266 34.727

153 3.629 34.875 153 4.798 35.030 154 1.748 34.668 154 2.253 34.726

154 3.649 34.878 154 4.742 35.024 155 1.688 34.665 155 2.244 34.725

155 3.661 34.879 155 4.727 35.022 156 1.655 34.662 156 2.253 34.726

156 3.675 34.883 156 4.709 35.020 157 1.629 34.661 157 2.269 34.728

157 3.752 34.908 157 4.696 35.018 158 1.630 34.660 158 2.318 34.734

158 4.147 34.956 158 4.691 35.017 159 1.638 34.661 159 2.332 34.736

159 4.131 34.951 159 4.654 35.013 160 1.631 34.661 160 2.267 34.733

160 4.091 34.947 160 4.620 35.011 161 1.615 34.661 161 2.210 34.729 Appendix A

Station 616 Station 632 Station 640 Station 641 Station 642

161 3.973 34.932 161 4.627 35.013 162 1.598 34.662 162 2.194 34.728

162 3.788 34.916 162 4.648 35.016 163 1.635 34.666 163 2.173 34.726

163 3.688 34.904 163 4.666 35.018 164 1.675 34.670 164 2.165 34.725

164 3.648 34.906 164 4.699 35.023 165 1.679 34.670 165 2.158 34.724

165 3.622 34.900 165 4.694 35.022 166 1.706 34.673 166 2.153 34.724

166 3.629 34.904 166 4.700 35.024 167 1.712 34.674 167 2.134 34.723

167 3.641 34.901 167 4.676 35.022 168 1.716 34.679 168 2.115 34.722

168 3.642 34.903 168 4.625 35.018 169 1.707 34.68 169 2.119 34.723

169 3.632 34.900 169 4.602 35.015 170 1.691 34.680 170 2.142 34.727

170 3.615 34.896 170 4.586 35.013 171 1.692 34.680 171 2.129 34.726

171 3.551 34.891 171 4.580 35.013 172 1.694 34.680 172 2.049 34.722

172 3.481 34.876 172 4.594 35.015 173 1.683 34.680 173 1.994 34.719

173 3.297 34.867 173 4.585 35.015 174 1.655 34.679 174 1.995 34.719

174 3.299 34.866 174 4.552 35.011 175 1.606 34.678 175 2.005 34.720

175 3.271 34.862 175 4.497 35.007 176 1.585 34.677 176 1.936 34.717

176 3.254 34.863 176 4.470 35.003 177 1.588 34.676 177 1.936 34.717

177 3.293 34.867 177 4.424 35.000 178 1.590 34.676 178 1.934 34.718

178 3.294 34.864 178 4.414 34.999 179 1.567 34.676 179 1.920 34.719

179 3.294 34.867 179 4.414 34.998 180 1.517 34.674 180 1.911 34.719

180 3.337 34.870 180 4.435 35.002 181 1.514 34.673 181 1.904 34.718

181 3.331 34.870 181 4.450 35.006 182 1.484 34.671 182 1.894 34.719

182 3.327 34.870 182 4.468 35.010 183 1.499 34.674 183 1.872 34.720

183 3.345 34.872 183 4.530 35.018 184 1.497 34.676 184 1.856 34.721

184 3.355 34.873 184 4.544 35.023 185 1.451 34.673 185 1.853 34.722

185 3.342 34.868 185 4.531 35.022 186 1.410 34.671 186 1.855 34.723

186 3.167 34.843 186 4.515 35.022 187 1.396 34.670 187 1.868 34.726

187 2.944 34.830 187 4.504 35.021 188 1.388 34.670 188 1.891 34.730

188 2.885 34.824 188 4.491 35.020 189 1.354 34.669 189 1.892 34.731

189 2.904 34.830 189 4.471 35.018 190 1.301 34.669 190 1.889 34.731

190 2.961 34.831 190 4.437 35.014 191 1.297 34.670 191 1.886 34.729

191 2.969 34.837 191 4.382 35.008 192 1.296 34.669 192 1.852 34.729

192 3.260 34.879 192 4.349 35.004 193 1.291 34.669 193 1.837 34.728

193 3.442 34.901 193 4.330 35.001 194 1.289 34.669 194 1.818 34.727

194 3.501 34.902 194 4.264 34.994 195 1.29 34.669 195 1.786 34.725

195 3.491 34.902 195 4.189 34.985 196 1.289 34.669 196 1.710 34.720

196 3.453 34.898 196 4.172 34.982 197 1.277 34.669 197 1.648 34.716

197 3.378 34.892 197 4.142 34.978 198 1.264 34.670 198 1.644 34.715

198 3.339 34.889 198 4.041 34.969 199 1.259 34.669 199 1.639 34.714

199 3.313 34.887 199 3.997 34.964 200 1.252 34.670 200 1.638 34.715

200 3.348 34.895 200 3.952 34.959 201 1.251 34.670 201 1.613 34.713

201 3.266 34.875 201 3.893 34.953 202 1.248 34.670 202 1.545 34.710

202 2.950 34.856 202 3.837 34.949 203 1.246 34.671 203 1.490 34.707

203 2.955 34.856 203 3.824 34.947 204 1.242 34.671 204 1.387 34.701

204 2.952 34.854 204 3.807 34.945 205 1.233 34.671 205 1.293 34.697

205 2.997 34.865 205 3.801 34.944 206 1.226 34.672 206 1.201 34.693

206 3.071 34.870 206 3.634 34.927 207 1.220 34.672 207 1.186 34.690

207 3.099 34.875 207 3.550 34.920 208 1.213 34.672 208 1.173 34.689

208 3.145 34.880 208 3.547 34.918 209 1.206 34.672 209 1.168 34.689

209 3.264 34.897 209 3.556 34.920 210 1.195 34.673 210 1.153 34.689

210 3.382 34.915 210 3.537 34.917 211 1.185 34.673 211 1.142 34.688

211 3.484 34.921 211 3.474 34.912 212 1.181 34.674 212 1.129 34.687

212 3.479 34.921 212 3.465 34.912 213 1.182 34.674 213 1.116 34.687

213 3.508 34.928 213 3.464 34.912 214 1.183 34.674 214 1.099 34.686

214 3.535 34.931 214 3.444 34.910 215 1.183 34.674 215 1.101 34.687

215 3.526 34.930 215 3.394 34.906 216 1.181 34.674 216 1.111 34.689

216 3.516 34.931 216 3.379 34.904 217 1.157 34.674 217 1.104 34.690 Appendix A

Station 616 Station 632 Station 640 Station 641 Station 642

217 3.528 34.934 217 3.368 34.903 218 1.145 34.675 218 1.093 34.69

218 3.593 34.945 218 3.366 34.903 219 1.132 34.676 219 1.091 34.690

219 3.657 34.952 219 3.406 34.909 220 1.125 34.676 220 1.087 34.690

220 3.602 34.945 220 3.533 34.927 221 1.118 34.679 221 1.079 34.692

221 3.519 34.941 221 3.533 34.928 222 1.111 34.683 222 1.082 34.693

222 3.465 34.937 222 3.520 34.927 223 1.108 34.684 223 1.085 34.695

223 3.390 34.930 223 3.505 34.927 224 1.098 34.687 224 1.075 34.695

224 3.357 34.927 224 3.504 34.927 225 1.084 34.692 225 1.082 34.697

225 3.325 34.922 225 3.432 34.922 226 1.074 34.694 226 1.102 34.7

226 3.226 34.911 226 3.343 34.917 227 1.062 34.698 227 1.087 34.703

227 3.143 34.905 227 3.339 34.917 228 1.060 34.700 228 1.096 34.708

228 3.061 34.899 228 3.331 34.916 229 1.050 34.707 229 1.086 34.711

229 3.042 34.9 229 3.286 34.913 230 1.043 34.713 230 1.080 34.712

230 3.040 34.899 230 3.216 34.909 231 1.068 34.698 231 1.096 34.718

231 3.038 34.899 231 3.129 34.901 232 1.114 34.721

232 3.037 34.898 232 3.044 34.893 233 1.096 34.722

233 3.016 34.898 233 2.963 34.886 234 1.083 34.722

234 3.010 34.900 234 2.879 34.878 235 1.074 34.722

235 3.006 34.898 235 2.849 34.873 236 1.080 34.723

236 2.946 34.890 236 2.850 34.873 237 1.079 34.723

237 2.886 34.887 237 2.815 34.874 238 1.080 34.723

238 2.842 34.884 238 2.746 34.871 239 1.077 34.723

239 2.761 34.869 239 2.676 34.866 240 1.089 34.726

240 2.577 34.859 240 2.613 34.862 241 1.096 34.728

241 2.540 34.856 241 2.576 34.861 242 1.095 34.730

242 2.513 34.853 242 2.561 34.863 243 1.092 34.731

243 2.475 34.847 243 2.562 34.866 244 1.170 34.740

244 2.424 34.843 244 2.562 34.869 245 1.269 34.750

245 2.315 34.833 245 2.550 34.872 246 1.224 34.751

246 2.243 34.832 246 2.528 34.874 247 1.143 34.751

247 2.233 34.831 247 2.507 34.873 248 1.110 34.751

248 2.221 34.830 248 2.424 34.870 249 1.119 34.755

249 2.219 34.830 249 2.346 34.864 250 1.120 34.758

250 2.197 34.827 250 2.323 34.865 251 1.120 34.76

251 2.159 34.824 251 2.233 34.863 252 1.123 34.762

252 2.096 34.820 252 2.207 34.862 253 1.111 34.764

253 2.058 34.819 253 2.201 34.861 254 1.102 34.764

254 2.046 34.818 254 2.198 34.861 255 1.112 34.765

255 2.033 34.817 255 2.199 34.861 256 1.124 34.768

256 2.019 34.816 256 2.202 34.863 257 1.128 34.770

257 2.014 34.816 257 2.182 34.865 258 1.138 34.773

258 2.018 34.816 258 2.133 34.864 259 1.096 34.773

259 2.019 34.816 259 2.115 34.863 260 1.084 34.774

260 1.996 34.814 260 2.110 34.863 261 1.094 34.777

261 1.998 34.816 261 2.100 34.863 262 1.100 34.779

262 1.999 34.816 262 2.090 34.862 263 1.098 34.782

263 1.995 34.816 263 2.076 34.862 264 1.072 34.783

264 1.990 34.816 264 2.063 34.861 265 1.070 34.784

265 1.981 34.816 265 2.013 34.859 266 1.073 34.786

266 1.954 34.813 266 1.974 34.858 267 1.065 34.788

267 1.917 34.813 267 1.976 34.859 268 1.048 34.79

268 1.913 34.812 268 1.958 34.859 269 0.994 34.792

269 1.870 34.812 269 1.900 34.857 270 0.961 34.792

270 1.862 34.813 270 1.867 34.854 271 0.945 34.792

271 1.865 34.813 271 1.857 34.854 272 0.950 34.793

272 1.858 34.813 272 1.836 34.854 273 0.962 34.797 Appendix A

Station 616 Station 632 Station 640 Station 641 Station 642

273 1.856 34.813 273 1.821 34.852 274 0.931 34.798

274 1.847 34.812 274 1.809 34.851 275 0.899 34.799

275 1.804 34.810 275 1.803 34.851 276 0.879 34.802

276 1.783 34.808 276 1.794 34.850 277 0.855 34.805

277 1.729 34.803 277 1.798 34.851 278 0.816 34.808

278 1.673 34.802 278 1.797 34.851 279 0.774 34.811

279 1.648 34.799 279 1.798 34.852 280 0.756 34.813

280 1.611 34.796 280 1.791 34.851 281 0.729 34.815

281 1.552 34.793 281 1.789 34.851 282 0.701 34.817

282 1.541 34.792 282 1.733 34.848 283 0.676 34.820

283 1.526 34.790 283 1.721 34.849 284 0.644 34.825

284 1.505 34.789 284 1.712 34.848 285 0.601 34.833

285 1.485 34.787 285 1.662 34.847 286 0.589 34.834

286 1.477 34.787 286 1.643 34.847 287 0.575 34.835

287 1.477 34.787 287 1.634 34.847 288 0.559 34.838

288 1.475 34.786 288 1.625 34.847 289 0.523 34.844

289 1.466 34.785 289 1.625 34.846 290 0.484 34.851

290 1.451 34.784 290 1.624 34.846 291 0.475 34.853

291 1.435 34.783 291 1.618 34.846 292 0.464 34.855

292 1.428 34.781 292 1.61 34.846 293 0.440 34.860

293 1.401 34.780 293 1.601 34.846 294 0.434 34.861

294 1.356 34.776 294 1.597 34.846 295 0.428 34.861

295 1.315 34.777 295 1.586 34.846 296 0.417 34.863

296 1.325 34.777 296 1.573 34.846 297 0.409 34.864

297 1.332 34.778 297 1.549 34.845 298 0.402 34.865

298 1.345 34.780 298 1.534 34.845 299 0.394 34.867

299 1.338 34.778 299 1.512 34.845 300 0.389 34.867

300 1.295 34.774 300 1.504 34.846 301 0.373 34.871

301 1.270 34.775 301 1.494 34.845 302 0.362 34.873

302 1.261 34.775 302 1.483 34.845 303 0.356 34.874

303 1.275 34.776 303 1.472 34.845 304 0.347 34.875

304 1.264 34.774 304 1.466 34.845 305 0.333 34.877

305 1.249 34.773 305 1.462 34.845 306 0.333 34.877

306 1.242 34.773 306 1.461 34.845 307 0.329 34.878

307 1.232 34.771 307 1.460 34.845 308 0.324 34.878

308 1.191 34.770 308 1.453 34.845 309 0.317 34.880

309 1.192 34.770 309 1.436 34.845 310 0.308 34.882

310 1.190 34.770 310 1.425 34.845 311 0.296 34.885

311 1.195 34.770 311 1.411 34.845 312 0.290 34.886

312 1.190 34.769 312 1.403 34.845 313 0.286 34.887

313 1.188 34.769 313 1.399 34.845 314 0.283 34.888

314 1.179 34.767 314 1.388 34.845 315 0.274 34.889

315 1.165 34.767 315 1.364 34.845 316 0.272 34.89

316 1.147 34.766 316 1.358 34.845 317 0.270 34.890

317 1.145 34.766 317 1.354 34.845 318 0.268 34.891

318 1.145 34.764 318 1.352 34.845 319 0.267 34.891

319 1.118 34.762 319 1.350 34.845 320 0.265 34.892

320 1.001 34.754 320 1.348 34.844 321 0.261 34.893

321 0.962 34.754 321 1.345 34.844 322 0.258 34.893

322 0.964 34.754 322 1.341 34.844 323 0.257 34.893

323 0.964 34.754 323 1.337 34.844 324 0.254 34.894

324 0.963 34.754 324 1.336 34.844 325 0.253 34.894

325 0.963 34.753 325 1.335 34.844 326 0.252 34.894

326 0.963 34.753 326 1.333 34.844 327 0.252 34.895

327 0.962 34.753 327 1.327 34.844 328 0.251 34.895

328 0.962 34.754 328 1.325 34.844 329 0.252 34.894 Appendix A

Station 616 Station 632 Station 640 Station 641 Station 642

329 0.960 34.753 329 1.324 34.844 330 0.251 34.895

330 0.953 34.754 330 1.322 34.844 331 0.249 34.895

331 0.955 34.754 331 1.318 34.843 332 0.249 34.895

332 0.954 34.753 332 1.319 34.844 333 0.248 34.895

333 0.950 34.753 334 0.246 34.896

334 0.932 34.753 335 0.244 34.897

335 0.924 34.755 336 0.241 34.897

336 0.925 34.755 337 0.240 34.898

337 0.926 34.755 338 0.238 34.898

338 0.928 34.755 339 0.238 34.898

339 0.926 34.755 340 0.235 34.900

340 0.929 34.755 341 0.237 34.899

341 0.936 34.756

342 0.943 34.757

343 0.945 34.758

344 0.947 34.759

345 0.947 34.759

346 0.947 34.759

347 0.946 34.759

348 0.940 34.760

349 0.932 34.760

350 0.931 34.760

351 0.929 34.760

352 0.928 34.760

353 0.928 34.760

354 0.928 34.760

355 0.925 34.760

356 0.923 34.761

357 0.923 34.761

358 0.924 34.761

359 0.923 34.761

360 0.923 34.761

361 0.924 34.762

362 0.925 34.762

363 0.928 34.762

364 0.930 34.763

365 0.928 34.763

366 0.923 34.763

367 0.913 34.763

368 0.905 34.764

369 0.907 34.764

370 0.902 34.765

371 0.900 34.765

372 0.886 34.765

373 0.880 34.766

374 0.880 34.766

375 0.879 34.766

376 0.875 34.765

377 0.872 34.767

378 0.864 34.766

379 0.858 34.767

380 0.856 34.767

381 0.854 34.767

382 0.854 34.767

383 0.852 34.767

384 0.851 34.767 Appendix A

Station 616 Station 632 Station 640 Station 641 Station 642

385 0.851 34.767

386 0.850 34.767

387 0.845 34.767

388 0.842 34.768

389 0.841 34.768

390 0.840 34.768

391 0.839 34.768

392 0.837 34.767

393 0.828 34.768

394 0.827 34.769

395 0.827 34.769

396 0.826 34.769

397 0.824 34.769

398 0.813 34.769

399 0.804 34.770

400 0.795 34.770

401 0.790 34.770

402 0.785 34.770

403 0.784 34.770

404 0.784 34.770

405 0.784 34.770

406 0.782 34.770

407 0.772 34.770

408 0.767 34.771

409 0.761 34.771

410 0.761 34.771

411 0.760 34.771

412 0.759 34.771

413 0.756 34.771

414 0.756 34.771

415 0.762 34.771 Appendix A

Station 643 Station 669 Station 699 Station 706

Depth Temperature Salinity Depth Temperature Salinity Depth Temperature Salinity Depth Temperature Salinity [m] [°C] (PSS) [m] [°C] (PSS) [m] [°C] (PSS) [m] [°C] (PSS)

3 5.705 32.192 3 6.749 32.879 3 5.834 33.984 3 5.495 34.073

4 5.777 32.475 4 6.720 32.980 4 5.834 34.002 4 5.491 34.073

5 5.817 32.609 5 6.692 33.040 5 5.837 33.955 5 5.493 34.087

6 5.813 32.725 6 6.662 33.098 6 5.867 34.108 6 5.492 34.089

7 5.846 32.817 7 6.623 33.157 7 5.946 34.376 7 5.5 34.112

8 5.868 32.872 8 6.592 33.201 8 5.929 34.443 8 5.522 34.154

9 5.869 33.037 9 6.564 33.237 9 5.868 34.452 9 5.603 34.276

10 5.888 33.232 10 6.539 33.264 10 5.802 34.453 10 5.764 34.468

11 6.006 33.386 11 6.519 33.286 11 5.687 34.457 11 5.711 34.479

12 6.06 33.441 12 6.494 33.316 12 5.524 34.470 12 5.617 34.482

13 6.080 33.454 13 6.473 33.344 13 5.415 34.474 13 5.522 34.485

14 5.994 33.482 14 6.447 33.383 14 5.289 34.485 14 5.430 34.490

15 5.961 33.588 15 6.433 33.407 15 5.257 34.482 15 5.430 34.510

16 5.946 33.705 16 6.416 33.449 16 5.238 34.485 16 5.404 34.526

17 5.836 33.784 17 6.406 33.506 17 5.324 34.510 17 5.360 34.536

18 5.832 33.854 18 6.395 33.633 18 5.400 34.527 18 5.345 34.534

19 5.777 33.895 19 6.405 33.792 19 5.403 34.533 19 5.33 34.533

20 5.745 33.912 20 6.416 33.885 20 5.409 34.537 20 5.318 34.532

21 5.695 33.917 21 6.409 34.005 21 5.467 34.557 21 5.319 34.532

22 5.669 33.924 22 6.358 34.079 22 5.499 34.567 22 5.317 34.532

23 5.636 33.939 23 6.307 34.152 23 5.442 34.566 23 5.300 34.532

24 5.611 33.956 24 6.317 34.162 24 5.382 34.562 24 5.292 34.531

25 5.607 33.967 25 6.307 34.186 25 5.321 34.559 25 5.287 34.531

26 5.766 34.022 26 6.277 34.266 26 5.365 34.592 26 5.290 34.531

27 5.905 34.083 27 6.239 34.344 27 5.356 34.609 27 5.283 34.531

28 5.890 34.102 28 6.200 34.428 28 5.340 34.617 28 5.268 34.531

29 5.796 34.118 29 6.165 34.502 29 5.277 34.617 29 5.263 34.531

30 5.678 34.120 30 6.130 34.571 30 5.132 34.617 30 5.257 34.531

31 5.455 34.106 31 6.093 34.639 31 5.09 34.616 31 5.238 34.533

32 4.691 34.058 32 6.045 34.747 32 5.022 34.611 32 5.224 34.538

33 4.067 34.017 33 6.053 34.793 33 4.942 34.607 33 5.205 34.541

34 4.067 34.009 34 6.073 34.806 34 4.852 34.599 34 5.169 34.540

35 3.964 34.035 35 6.128 34.840 35 4.713 34.593 35 5.096 34.54

36 3.989 34.082 36 6.164 34.866 36 4.599 34.583 36 5.057 34.539

37 4.082 34.128 37 6.167 34.892 37 4.500 34.575 37 4.983 34.535

38 4.025 34.146 38 6.240 34.921 38 4.395 34.567 38 4.896 34.532

39 3.815 34.146 39 6.278 34.950 39 4.369 34.563 39 4.895 34.536

40 4.089 34.205 40 6.301 34.970 40 4.346 34.560 40 4.812 34.535

41 4.736 34.355 41 6.310 34.975 41 4.336 34.558 41 4.694 34.531

42 5.100 34.424 42 6.313 34.981 42 4.316 34.555 42 4.660 34.527

43 5.060 34.434 43 6.313 34.983 43 4.295 34.553 43 4.65 34.527

44 4.850 34.452 44 6.323 34.987 44 4.284 34.552 44 4.648 34.526

45 5.110 34.532 45 6.324 34.989 45 4.270 34.551 45 4.630 34.525

46 4.953 34.545 46 6.324 34.991 46 4.212 34.544 46 4.618 34.524

47 4.929 34.580 47 6.325 34.996 47 4.144 34.549 47 4.602 34.524

48 5.100 34.631 48 6.307 35.004 48 4.173 34.556 48 4.526 34.520

49 5.258 34.681 49 6.298 35.010 49 4.150 34.559 49 4.495 34.522

50 5.089 34.673 50 6.299 35.012 50 4.132 34.561 50 4.466 34.524

51 4.614 34.646 51 6.271 35.012 51 4.019 34.558 51 4.457 34.528

52 4.257 34.624 52 6.244 35.009 52 3.947 34.558 52 4.396 34.530

53 4.229 34.617 53 6.119 35.000 53 3.898 34.566 53 4.394 34.532

54 4.281 34.625 54 5.997 34.992 54 3.885 34.566 54 4.403 34.535

55 4.378 34.644 55 5.979 34.989 55 3.874 34.568 55 4.403 34.541

56 4.407 34.654 56 5.909 34.982 56 3.864 34.572 56 4.374 34.546

57 4.104 34.638 57 5.829 34.974 57 3.856 34.577 57 4.315 34.544

58 3.718 34.606 58 5.553 34.952 58 3.842 34.578 58 4.060 34.542 Appendix A

Station 643 Station 669 Station 699 Station 706

Depth Temperature Salinity Depth Temperature Salinity Depth Temperature Salinity Depth Temperature Salinity [m] [°C] (PSS) [m] [°C] (PSS) [m] [°C] (PSS) [m] [°C] (PSS)

59 3.685 34.599 59 5.256 34.924 59 3.830 34.578 59 3.897 34.549

60 4.126 34.667 60 5.135 34.911 60 3.814 34.579 60 3.871 34.552

61 4.135 34.670 61 5.095 34.903 61 3.797 34.585 61 3.811 34.554

62 3.934 34.660 62 4.984 34.895 62 3.816 34.59 62 3.781 34.557

63 3.902 34.668 63 4.993 34.9 63 3.787 34.589 63 3.749 34.560

64 3.878 34.681 64 4.911 34.904 64 3.753 34.591 64 3.736 34.559

65 3.859 34.690 65 4.841 34.906 65 3.692 34.593 65 3.733 34.558

66 3.809 34.687 66 4.844 34.904 66 3.656 34.597 66 3.727 34.560

67 3.824 34.691 67 4.820 34.902 67 3.646 34.597 67 3.718 34.565

68 3.835 34.695 68 4.784 34.901 68 3.636 34.598 68 3.716 34.571

69 3.824 34.695 69 4.775 34.9 69 3.628 34.598

70 3.821 34.696 70 4.781 34.902 70 3.586 34.597

71 3.802 34.695 71 4.785 34.907 71 3.556 34.598

72 3.789 34.694 72 4.773 34.914 72 3.554 34.598

73 3.778 34.694 73 4.726 34.914 73 3.556 34.597

74 3.801 34.702 74 4.734 34.916 74 3.538 34.600

75 3.821 34.709 75 4.739 34.922 75 3.537 34.600

76 3.898 34.732 76 4.701 34.921 76 3.536 34.600

77 3.990 34.753 77 4.706 34.922 77 3.535 34.601

78 3.940 34.752 78 4.712 34.924 78 3.544 34.603

79 3.878 34.749 79 4.680 34.922 79 3.561 34.605

80 3.825 34.746 80 4.649 34.921 80 3.570 34.610

81 3.744 34.740 81 4.574 34.917 81 3.624 34.628

82 3.708 34.737 82 4.493 34.915 82 3.651 34.644

83 3.681 34.735 83 4.306 34.911 83 3.776 34.664

84 3.632 34.731 84 4.290 34.908 84 3.816 34.669

85 3.584 34.728 85 4.288 34.906 85 3.810 34.670

86 3.542 34.727 86 4.241 34.905 86 3.815 34.672

87 3.550 34.728 87 4.227 34.901 87 3.782 34.670

88 3.501 34.727 88 4.171 34.899 88 3.795 34.683

89 3.517 34.730 89 4.149 34.896 89 3.804 34.685

90 3.569 34.745 90 4.129 34.895 90 3.820 34.688

91 3.500 34.746 91 4.083 34.896 91 3.825 34.696

92 3.432 34.742 92 4.019 34.906 92 3.830 34.698

93 3.434 34.741 93 4.006 34.909 93 3.829 34.698

94 3.687 34.779 94 3.980 34.909 94 3.810 34.698

95 3.728 34.787 95 4.051 34.917 95 3.722 34.697

96 3.643 34.781 96 4.065 34.919 96 3.540 34.703

97 3.586 34.775 97 4.061 34.918 97 3.460 34.702

98 3.463 34.766 98 4.066 34.919 98 3.452 34.703

99 3.464 34.772 99 3.898 34.899 99 3.421 34.704

100 3.209 34.753 100 3.814 34.891 100 3.377 34.705

101 3.102 34.741 101 3.804 34.888 101 3.311 34.706

102 3.124 34.741 102 3.8 34.887 102 3.277 34.708

103 3.270 34.758 103 3.803 34.887 103 3.257 34.708

104 3.407 34.774 104 3.853 34.891 104 3.234 34.709

105 3.416 34.776 105 3.836 34.888 105 3.212 34.709

106 3.437 34.779 106 3.789 34.885 106 3.194 34.710

107 3.456 34.782 107 3.789 34.885 107 3.171 34.708

108 3.497 34.788 108 3.804 34.887 108 3.062 34.714

109 3.642 34.808 109 3.820 34.889 109 3.010 34.718

110 3.680 34.815 110 3.881 34.896 110 3.001 34.718

111 3.650 34.813 111 3.809 34.887 111 2.997 34.718

112 3.470 34.800 112 3.875 34.896 112 2.995 34.718

113 3.263 34.785 113 3.906 34.899 113 2.991 34.718

114 3.226 34.779 114 3.901 34.898 114 2.987 34.717 Appendix A

Station 643 Station 669 Station 699 Station 706

Depth Temperature Salinity Depth Temperature Salinity Depth Temperature Salinity Depth Temperature Salinity [m] [°C] (PSS) [m] [°C] (PSS) [m] [°C] (PSS) [m] [°C] (PSS)

115 3.121 34.771 115 3.787 34.888 115 2.976 34.717

116 3.065 34.765 116 3.874 34.896 116 2.955 34.718

117 3.029 34.761 117 3.871 34.894 117 2.924 34.718

118 2.982 34.757 118 3.839 34.892 118 2.889 34.723

119 2.951 34.753 119 3.795 34.888 119 2.883 34.724

120 2.937 34.751 120 3.750 34.879 120 2.863 34.726

121 2.885 34.747 121 3.726 34.876 121 2.855 34.727

122 2.858 34.745 122 3.723 34.877 122 2.851 34.727

123 2.816 34.742 123 3.736 34.880 123 2.845 34.727

124 2.782 34.739 124 3.734 34.88 124 2.842 34.727

125 2.780 34.739 125 3.730 34.879 125 2.808 34.721

126 3.188 34.789 126 3.724 34.878 126 2.778 34.731

127 3.513 34.831 127 3.717 34.877 127 2.769 34.732

128 3.495 34.834 128 3.720 34.879 128 2.771 34.731

129 3.463 34.834 129 3.729 34.880 129 2.768 34.731

130 3.459 34.835 130 3.731 34.881 130 2.770 34.730

131 3.451 34.835 131 3.748 34.881

132 3.432 34.834 132 3.728 34.879

133 3.384 34.829 133 3.721 34.879

134 3.252 34.820 134 3.737 34.881

135 3.251 34.818 135 3.755 34.885

136 3.25 34.817 136 3.812 34.895

137 3.252 34.817 137 3.827 34.898

138 3.299 34.823 138 3.838 34.901

139 3.442 34.841 139 3.836 34.898

140 3.451 34.845 140 3.773 34.889

141 3.389 34.841 141 3.767 34.889

142 3.239 34.830 142 3.779 34.891

143 3.173 34.825 143 3.783 34.893

144 3.075 34.818 144 3.798 34.895

145 3.063 34.816 145 3.819 34.898

146 3.044 34.814 146 3.818 34.898

147 3.041 34.814 147 3.817 34.900

148 3.040 34.814

149 3.029 34.813

150 3.001 34.812

151 2.966 34.809

152 2.976 34.811

153 2.978 34.812

154 2.985 34.815

155 2.975 34.816

156 2.994 34.818

157 2.949 34.816

158 2.896 34.813

159 2.947 34.818

160 2.978 34.822

161 2.983 34.824

162 2.937 34.821

163 2.847 34.815

164 2.795 34.810

165 2.765 34.807

166 2.713 34.803

167 2.670 34.799

168 2.603 34.794

169 2.527 34.788

170 2.526 34.787 Appendix A

Station 643 Station 669 Station 699 Station 706

Depth Temperature Salinity Depth Temperature Salinity Depth Temperature Salinity Depth Temperature Salinity [m] [°C] (PSS) [m] [°C] (PSS) [m] [°C] (PSS) [m] [°C] (PSS)

171 2.525 34.786

172 2.525 34.785

173 2.411 34.777

174 2.345 34.771

175 2.310 34.767

176 2.242 34.761

177 2.239 34.759

178 2.244 34.759

179 2.252 34.758

180 2.257 34.760

181 2.246 34.758

182 2.116 34.748

183 2.080 34.744

184 2.075 34.743

185 2.057 34.741

186 2.042 34.740

187 2.028 34.739

188 2.007 34.737

189 2.003 34.737

190 2.019 34.738

191 1.996 34.737

192 1.962 34.735

193 1.951 34.734

194 1.898 34.732

195 1.848 34.728

196 1.836 34.727

197 1.830 34.727

198 1.834 34.727

199 1.829 34.729

200 1.844 34.731

201 1.829 34.731

202 1.822 34.730

203 1.830 34.733

204 1.819 34.733

205 1.830 34.737

206 1.823 34.737

207 1.809 34.736

208 1.778 34.734

209 1.697 34.731

210 1.642 34.729

211 1.611 34.726

212 1.572 34.723

213 1.522 34.720

214 1.511 34.719

215 1.502 34.718

216 1.510 34.718

217 1.552 34.722

218 1.555 34.723

219 1.560 34.725

220 1.573 34.727

221 1.596 34.730

222 1.555 34.729

223 1.339 34.718

224 1.298 34.715

225 1.303 34.715

226 1.314 34.716 Appendix A

Station 643 Station 669 Station 699 Station 706

Depth Temperature Salinity Depth Temperature Salinity Depth Temperature Salinity Depth Temperature Salinity [m] [°C] (PSS) [m] [°C] (PSS) [m] [°C] (PSS) [m] [°C] (PSS)

227 1.277 34.715

228 1.232 34.713

229 1.227 34.712

230 1.224 34.712

231 1.209 34.711

232 1.214 34.711

233 1.216 34.712

234 1.241 34.716

235 1.259 34.719

236 1.244 34.718

237 1.234 34.717

238 1.235 34.718

239 1.249 34.720

240 1.294 34.725

241 1.356 34.732

242 1.367 34.737

243 1.393 34.749

244 1.353 34.754

245 1.354 34.758

246 1.349 34.762

247 1.347 34.764

248 1.341 34.765

249 1.327 34.765

250 1.325 34.767

251 1.334 34.768

252 1.337 34.768

253 1.343 34.773

254 1.353 34.776

255 1.351 34.776

256 1.355 34.777

257 1.38 34.784

258 1.406 34.790

259 1.385 34.789

260 1.339 34.788

261 1.318 34.788

262 1.327 34.788

263 1.342 34.790

264 1.347 34.790

265 1.349 34.791

266 1.352 34.792

267 1.351 34.792

268 1.347 34.793

269 1.321 34.794

270 1.325 34.793

271 1.327 34.793

272 1.322 34.794

273 1.312 34.794

274 1.304 34.794

275 1.255 34.796

276 1.144 34.801

277 1.2 34.791 Appendix B

Appendix B

Raw data PAR-measurements [Note: Data collected by M. Wisshak ([email protected]) during MSM 02/03 expedition of RV Maria S. Merian in 2006]

Station 654 Station 672/1 Station 672/2

depth PAR % surface depth PAR % surface depth PAR % surface [m] [µmol photons m2 s1] illumination [m] [µmol photons m2 s1] illumination [m] [µmol photons m2 s1] illumination

air 635.70 air 1066.81 air 2579.70

0 276.20 100.0 0 393.37 100.0 0 1051.83 100.0

1 231.92 84.0 1 337.78 85.9 1 455.19 43.3

2.5 184.92 67.0 2.5 263.04 66.9 2.5 351.70 33.4

5 132.44 48.0 5 162.73 41.4 5 203.81 19.4

7.5 94.71 34.3 7.5 101.99 25.9 7.5 133.97 12.7

10 39.63 14.3 10 61.55 15.6 10 80.10 7.6

15 19.32 7.0 15 22.68 5.8 15 29.74 2.8

20 12.84 4.6 20 8.78 2.2 20 11.92 1.1

25 8.26 3.0 25 3.37 0.9 25 4.77 0.5

30 6.54 2.4 30 1.55 0.4 30 2.24 0.2

35 4.00 1.4 35 0.77 0.2 35 1.30 0.1

40 3.32 1.2 40 0.36 0.1 40 0.66 0.1

45 2.16 0.8 45 0.14 0.0 45 0.43 0.0

50 1.29 0.5 50 0.06 0.0 50 0.33 0.0

55 0.73 0.3 55 55 0.23 0.0

60 0.74 0.3 60 60 0.24 0.0

65 0.44 0.2 65 65 0.06 0.0

70 0.41 0.1 70 70

75 0.38 0.1 75 75

Station 702 Station 712

depth PAR % surface depth PAR % surface [m] [µmol photons m2 s1] illumination [m] [µmol photons m2 s1] illumination

air 878.95 air 729.92

0 270.27 100.0 0 217.96 100.00

1 172.54 63.8 1 206.69 94.83

2.5 160.91 59.5 2.5 165.92 76.12

5 152.86 56.6 5 129.51 59.42

7.5 117.78 43.6 7.5 94.85 43.52

10 87.06 32.2 10 69.31 31.80

15 45.81 17.0 15 37.22 17.08

20 26.38 9.8 20 21.64 9.93

25 16.10 6.0 25 12.88 5.91

30 10.18 3.8 30 7.93 3.64

35 5.85 2.2 35 4.99 2.29

40 3.70 1.4 40 3.27 1.50

45 2.16 0.8 45 2.06 0.94

50 1.58 0.6 50 1.34 0.61

55 0.89 0.3 55 0.88 0.41

60 0.49 0.2 60 0.67 0.31

65 65 0.45 0.21

70 70 0.25 0.12

75 75 0.12 0.06 Appendix C

Appendix C

Raw data rhodolith size- and morphology-measurements

Rhodolith # Site Long axis (L) Intermediate axis (I) Short axis (S) Sphericity Size [cm] [cm] [cm] [(S2/LI)0.5] [(LIS/4π)0.5]

R_637_01 Krossfjorden 7.11 2.85 1.93 0.57 11.03

R_637_02 Krossfjorden 6.33 3.30 2.90 0.74 17.09

R_637_03 Krossfjorden 4.49 4.33 3.39 0.84 18.59

R_637_04 Krossfjorden 6.39 3.48 3.26 0.78 20.45

R_637_05 Krossfjorden 4.91 3.04 2.61 0.77 10.99

R_637_06 Krossfjorden 5.86 3.92 3.68 0.84 23.85

R_637_07 Krossfjorden 4.11 4.02 2.77 0.77 12.91

R_637_08 Krossfjorden 7.07 2.83 1.43 0.47 8.07

R_637_09 Krossfjorden 5.14 4.08 3.51 0.84 20.76

R_637_10 Krossfjorden 5.06 3.22 2.03 0.63 9.33

R_637_11 Krossfjorden 4.90 3.29 1.48 0.51 6.73

R_637_12 Krossfjorden 4.68 3.19 2.14 0.67 9.01

R_637_13 Krossfjorden 5.42 2.46 1.16 0.47 4.36

R_637_14 Krossfjorden 5.01 3.25 2.23 0.67 10.24

R_637_15 Krossfjorden 4.62 3.34 2.18 0.68 9.49

R_637_16 Krossfjorden 4.56 4.24 3.20 0.81 17.45

R_637_17 Krossfjorden 4.88 3.19 1.85 0.60 8.12

R_637_18 Krossfjorden 4.39 2.65 2.34 0.78 7.68

R_637_19 Krossfjorden 4.52 3.48 2.41 0.72 10.69

R_637_20 Krossfjorden 4.55 2.79 1.35 0.52 4.83

R_637_21 Krossfjorden 4.67 2.44 1.78 0.65 5.72

R_637_22 Krossfjorden 5.21 2.93 1.73 0.58 7.45

R_637_23 Krossfjorden 3.73 2.60 2.40 0.84 6.57

R_637_24 Krossfjorden 3.56 3.32 2.40 0.79 8.00

R_637_25 Krossfjorden 3.92 2.91 2.89 0.90 9.30

R_637_26 Krossfjorden 4.31 2.13 0.89 0.44 2.30

R_637_27 Krossfjorden 3.49 2.35 2.13 0.82 4.93

R_637_28 Krossfjorden 2.46 3.00 2.46 0.94 5.12

R_637_29 Krossfjorden 4.03 3.27 1.82 0.63 6.77

R_637_30 Krossfjorden 4.14 2.33 1.69 0.67 4.60

R_637_31 Krossfjorden 4.13 2.83 2.13 0.73 7.02

R_637_32 Krossfjorden 3.58 3.17 2.40 0.80 7.68

R_637_33 Krossfjorden 3.99 3.20 1.31 0.51 4.72

R_637_34 Krossfjorden 4.07 2.38 0.88 0.43 2.40

R_637_35 Krossfjorden 3.96 2.93 1.80 0.65 5.89

R_637_36 Krossfjorden 3.62 2.12 1.30 0.60 2.81

R_637_37 Krossfjorden 3.64 3.11 2.18 0.75 6.96

R_637_38 Krossfjorden 4.11 2.09 1.06 0.51 2.57

R_637_39 Krossfjorden 3.97 2.08 1.19 0.56 2.77

R_637_40 Krossfjorden 3.99 1.83 1.22 0.59 2.51

R_637_41 Krossfjorden 3.76 2.42 1.45 0.61 3.72

R_637_42 Krossfjorden 3.87 2.47 1.44 0.60 3.88

R_637_43 Krossfjorden 3.29 2.90 2.06 0.76 5.54

R_637_44 Krossfjorden 3.61 2.93 2.63 0.87 7.85

R_637_45 Krossfjorden 3.72 2.49 2.21 0.81 5.77 Appendix C

Rhodolith # Site Long axis (L) Intermediate axis (I) Short axis (S) Sphericity Size [cm] [cm] [cm] [(S2/LI)0.5] [(LIS/4π)0.5]

R_637_46 Krossfjorden 3.56 2.73 2.16 0.78 5.92

R_637_47 Krossfjorden 3.49 2.86 1.69 0.66 4.76

R_637_48 Krossfjorden 4.34 1.99 1.49 0.64 3.63

R_637_49 Krossfjorden 4.25 2.44 1.37 0.57 4.01

R_637_50 Krossfjorden 3.42 2.78 2.36 0.84 6.33

R_637_51 Krossfjorden 3.21 2.68 1.75 0.71 4.25

R_637_52 Krossfjorden 3.49 3.09 1.89 0.69 5.75

R_637_53 Krossfjorden 3.12 3.02 2.19 0.80 5.82

R_637_54 Krossfjorden 3.46 2.60 0.84 0.43 2.13

R_637_55 Krossfjorden 2.92 2.45 2.23 0.89 4.50

R_637_56 Krossfjorden 3.76 1.40 0.99 0.57 1.47

R_637_57 Krossfjorden 3.10 2.56 2.20 0.85 4.93

R_637_58 Krossfjorden 3.29 2.22 1.79 0.76 3.69

R_637_59 Krossfjorden 3.00 2.53 1.98 0.80 4.24

R_637_60 Krossfjorden 3.17 2.36 1.82 0.76 3.84

R_637_61 Krossfjorden 3.31 2.92 1.25 0.54 3.41

R_637_62 Krossfjorden 3.00 2.32 1.66 0.73 3.26

R_637_63 Krossfjorden 3.12 2.17 1.42 0.67 2.71

R_637_64 Krossfjorden 2.37 1.46 0.83 0.58 0.81

R_637_65 Krossfjorden 3.50 1.42 1.20 0.66 1.68

R_637_66 Krossfjorden 3.18 2.29 1.31 0.62 2.69

R_637_67 Krossfjorden 3.29 1.92 1.36 0.66 2.42

R_637_68 Krossfjorden 2.62 1.95 1.14 0.63 1.64

R_637_69 Krossfjorden 2.69 1.68 1.57 0.82 2.00

R_637_70 Krossfjorden 2.86 1.75 1.47 0.76 2.08

R_637_71 Krossfjorden 3.01 2.53 1.31 0.61 2.81

R_637_72 Krossfjorden 2.93 2.17 1.43 0.69 2.56

R_637_73 Krossfjorden 3.29 2.19 1.61 0.71 3.27

R_637_74 Krossfjorden 2.59 2.20 1.83 0.84 2.94

R_637_75 Krossfjorden 3.15 2.46 2.08 0.82 4.55

R_637_76 Krossfjorden 2.93 2.13 1.77 0.79 3.12

R_637_77 Krossfjorden 2.34 2.21 1.89 0.88 2.76

R_637_78 Krossfjorden 2.96 2.06 1.87 0.83 3.22

R_637_79 Krossfjorden 2.58 2.35 2.01 0.87 3.44

R_637_80 Krossfjorden 2.56 2.09 1.67 0.80 2.52

R_637_81 Krossfjorden 2.78 1.35 1.20 0.73 1.27

R_637_82 Krossfjorden 3.08 2.04 0.88 0.50 1.56

R_637_83 Krossfjorden 2.48 2.09 1.22 0.66 1.78

R_637_84 Krossfjorden 3.04 2.24 1.59 0.72 3.05

R_637_85 Krossfjorden 2.53 1.78 1.57 0.82 1.99

R_637_86 Krossfjorden 2.66 2.04 1.06 0.59 1.62

R_637_87 Krossfjorden 2.54 2.34 2.16 0.92 3.62

R_637_88 Krossfjorden 2.64 1.71 1.40 0.76 1.78

R_637_89 Krossfjorden 2.15 1.41 1.21 0.78 1.03

R_637_90 Krossfjorden 3.13 2.55 1.25 0.58 2.81

R_637_91 Krossfjorden 2.20 1.72 0.97 0.63 1.04

R_637_92 Krossfjorden 2.71 2.11 1.50 0.73 2.42

R_637_93 Krossfjorden 2.80 2.10 1.75 0.80 2.90

R_637_94 Krossfjorden 2.76 1.74 1.29 0.70 1.75

R_637_95 Krossfjorden 3.06 1.70 1.41 0.73 2.07

R_637_96 Krossfjorden 2.70 2.57 1.75 0.76 3.43

R_637_97 Krossfjorden 2.27 2.06 1.25 0.69 1.65

R_637_98 Krossfjorden 2.71 1.86 0.89 0.54 1.27 Appendix C

Rhodolith # Site Long axis (L) Intermediate axis (I) Short axis (S) Sphericity Size [cm] [cm] [cm] [(S2/LI)0.5] [(LIS/4π)0.5]

R_637_99 Krossfjorden 2.96 2.44 1.01 0.52 2.06

R_637_100 Krossfjorden 2.80 2.10 1.43 0.70 2.37

R_637_101 Krossfjorden 2.54 2.10 1.89 0.87 2.84

R_637_102 Krossfjorden 2.58 1.42 0.85 0.58 0.88

R_637_103 Krossfjorden 2.68 2.25 1.49 0.72 2.53

R_637_104 Krossfjorden 2.39 1.25 1.21 0.79 1.02

R_637_105 Krossfjorden 2.59 1.38 0.79 0.56 0.80

R_644_01 Krossfjorden 5.60 4.40 3.50 0.79 24.33

R_644_02 Krossfjorden 4.37 3.54 2.87 0.81 12.52

R_644_03 Krossfjorden 4.31 3.41 3.14 0.88 13.02

R_644_04 Krossfjorden 6.59 3.46 2.95 0.73 18.97

R_644_05 Krossfjorden 5.66 5.12 3.93 0.81 32.13

R_644_06 Krossfjorden 6.97 3.21 2.86 0.72 18.05

R_644_07 Krossfjorden 4.33 4.21 2.54 0.71 13.06

R_644_08 Krossfjorden 5.65 3.81 3.34 0.80 20.28

R_644_09 Krossfjorden 3.99 3.09 2.09 0.71 7.27

R_644_10 Krossfjorden 6.44 4.74 3.33 0.71 28.68

R_644_11 Krossfjorden 4.86 3.68 3.12 0.82 15.74

R_644_12 Krossfjorden 5.39 4.19 3.11 0.75 19.81

R_644_13 Krossfjorden 4.85 3.93 2.12 0.62 11.40

R_644_14 Krossfjorden 6.16 4.03 3.99 0.86 27.94

R_644_15 Krossfjorden 8.76 6.01 5.12 0.79 76.04

R_652_01 Krossfjorden 7.90 5.00 3.10 0.62 34.54

R_652_02 Krossfjorden 5.20 3.80 2.60 0.70 14.49

R_671_01 Mosselbukta 11.59 9.34 8.22 0.85 251.01

R_671_02 Mosselbukta 13.02 11.15 8.00 0.76 327.62

R_671_03 Mosselbukta 12.00 8.92 8.14 0.85 245.79

R_671_04 Mosselbukta 11.99 8.50 6.11 0.72 175.66

R_671_05 Mosselbukta 14.00 10.83 8.78 0.80 375.53

R_671_06 Mosselbukta 9.94 9.41 8.68 0.93 229.03

R_671_07 Mosselbukta 4.05 2.99 2.23 0.74 7.62

R_671_09 Mosselbukta 4.60 4.14 2.58 0.70 13.86

R_671_10 Mosselbukta 6.07 4.50 4.43 0.90 34.14

R_671_11 Mosselbukta 17.20 14.52 7.90 0.63 556.57

R_671_12 Mosselbukta 16.20 12.18 7.20 0.64 400.77

R_671_13 Mosselbukta 12.16 10.60 7.66 0.77 278.52

R_671_14 Mosselbukta 14.38 11.10 8.05 0.74 362.47

R_671_15 Mosselbukta 5.24 3.00 1.85 0.60 8.20

R_671_16 Mosselbukta 4.80 2.87 1.97 0.66 7.66

R_681_01 Mosselbukta 11.87 9.44 7.14 0.77 225.69

R_682_01 Mosselbukta 14.47 14.00 9.13 0.74 521.75

R_684_01 Mosselbukta 8.47 7.97 6.79 0.88 129.30

R_684_02 Mosselbukta 14.25 12.30 8.04 0.72 397.53

R_684_03 Mosselbukta 9.11 5.16 3.46 0.63 45.88

R_684_04 Mosselbukta 10.85 6.20 4.80 0.70 91.09

R_684_05 Mosselbukta 10.66 7.68 4.81 0.66 111.09

R_684_06 Mosselbukta 10.19 9.33 6.20 0.74 166.28

R_684_07 Mosselbukta 13.22 8.13 7.55 0.81 228.91

R_684_08 Mosselbukta 10.78 9.62 7.32 0.80 214.14

R_684_09 Mosselbukta 10.99 10.28 5.96 0.68 189.95

R_684_10 Mosselbukta 6.20 4.77 3.87 0.80 32.29

R_684_11 Mosselbukta 6.25 4.97 1.07 0.33 9.38

R_684_12 Mosselbukta 6.79 4.24 1.52 0.43 12.34 Appendix C

Rhodolith # Site Long axis (L) Intermediate axis (I) Short axis (S) Sphericity Size [cm] [cm] [cm] [(S2/LI)0.5] [(LIS/4π)0.5]

R_684_13 Mosselbukta 3.00 2.76 1.00 0.49 2.34

R_684_14 Mosselbukta 9.11 7.12 6.72 0.89 122.96

R_684_15 Mosselbukta 10.33 7.63 5.41 0.72 120.29

R_684_16 Mosselbukta 12.59 8.70 6.52 0.73 201.46

R_684_17 Mosselbukta 10.87 8.52 8.16 0.90 213.18

R_684_18 Mosselbukta 7.93 6.27 5.12 0.81 71.81

R_684_19 Mosselbukta 8.60 7.95 6.80 0.88 131.15

R_684_20 Mosselbukta 8.99 8.41 6.68 0.84 142.47

R_684_21 Mosselbukta 10.02 8.51 5.64 0.72 135.67

R_684_22 Mosselbukta 15.25 10.27 5.72 0.59 252.72

R_684_23 Mosselbukta 12.93 9.73 6.26 0.68 222.17

R_684_24 Mosselbukta 9.77 8.52 7.08 0.84 166.25

R_684_25 Mosselbukta 5.47 3.43 1.97 0.59 10.43

R_684_26 Mosselbukta 5.61 5.46 2.63 0.61 22.73

R_684_27 Mosselbukta 7.49 5.53 3.18 0.63 37.16

R_684_28 Mosselbukta 6.76 5.14 4.75 0.87 46.56

R_684_29 Mosselbukta 8.20 6.72 5.60 0.83 87.05

R_684_30 Mosselbukta 8.68 6.27 2.19 0.45 33.62

R_684_31 Mosselbukta 10.43 9.55 5.69 0.69 159.88

R_684_32 Mosselbukta 7.82 5.65 1.57 0.38 19.57

R_684_33 Mosselbukta 11.11 7.72 7.06 0.83 170.82

R_684_34 Mosselbukta 8.80 8.89 5.85 0.76 129.10

R_684_35 Mosselbukta 7.59 7.37 6.75 0.93 106.51

R_684_36 Mosselbukta 9.82 7.13 6.85 0.88 135.30

R_684_38 Mosselbukta 7.21 6.24 2.30 0.49 29.19

R_684_39 Mosselbukta 11.24 8.87 8.66 0.91 243.56

R_684_40 Mosselbukta 10.72 8.05 7.72 0.88 187.93

R_684_41 Mosselbukta 9.24 7.77 4.73 0.68 95.80

R_684_42 Mosselbukta 11.54 8.27 2.55 0.41 68.65

R_684_43 Mosselbukta 9.56 8.89 5.61 0.72 134.50

R_684_44 Mosselbukta 11.88 11.11 6.48 0.68 241.27

R_684_45 Mosselbukta 12.29 10.35 8.55 0.83 306.80

R_684_46 Mosselbukta 9.56 8.13 6.68 0.83 146.46

R_684_47 Mosselbukta 11.15 7.02 5.96 0.77 131.60

R_684_48 Mosselbukta 12.02 8.09 7.34 0.82 201.35

R_684_49 Mosselbukta 9.07 6.77 4.61 0.70 79.85

R_684_50 Mosselbukta 8.07 7.06 6.71 0.92 107.84

R_684_51 Mosselbukta 11.56 8.96 6.72 0.76 196.35

R_684_52 Mosselbukta 11.19 9.00 8.06 0.86 228.98

R_684_53 Mosselbukta 7.60 6.09 5.45 0.86 71.16

R_684_54 Mosselbukta 8.22 5.61 3.91 0.69 50.86

R_684_55 Mosselbukta 7.79 6.56 5.83 0.87 84.04

R_684_56 Mosselbukta 9.12 8.07 6.09 0.80 126.44

R_684_57 Mosselbukta 13.68 10.02 6.82 0.70 263.71

R_684_58 Mosselbukta 9.57 8.10 6.52 0.82 142.57

R_684_59 Mosselbukta 11.36 9.90 6.11 0.69 193.84

R_684_60 Mosselbukta 6.88 4.44 4.15 0.83 35.76

R_684_61 Mosselbukta 4.86 4.05 1.81 0.55 10.05

R_684_62 Mosselbukta 9.60 6.25 6.65 0.90 112.56

R_684_63 Mosselbukta 9.86 8.93 6.28 0.77 155.99

R_684_64 Mosselbukta 10.39 8.05 6.83 0.82 161.15

R_684_65 Mosselbukta 8.15 7.72 5.92 0.82 105.07

R_684_66 Mosselbukta 8.77 8.54 6.02 0.79 127.19 Appendix C

Rhodolith # Site Long axis (L) Intermediate axis (I) Short axis (S) Sphericity Size [cm] [cm] [cm] [(S2/LI)0.5] [(LIS/4π)0.5]

R_684_67 Mosselbukta 11.10 8.64 6.26 0.74 169.36

R_684_68 Mosselbukta 10.93 8.98 6.49 0.75 179.70

R_684_69 Mosselbukta 14.78 10.21 8.48 0.78 360.99

R_684_70 Mosselbukta 12.50 6.33 5.36 0.71 119.64

R_684_71 Mosselbukta 10.17 7.92 5.28 0.70 119.97

R_684_72 Mosselbukta 8.24 5.45 5.80 0.91 73.48

R_684_73 Mosselbukta 9.21 7.53 6.20 0.82 121.29

R_684_74 Mosselbukta 9.39 8.05 6.25 0.80 133.27

R_684_75 Mosselbukta 10.78 7.54 6.25 0.78 143.31

R_684_76 Mosselbukta 8.19 8.71 7.20 0.90 144.89

R_684_77 Mosselbukta 10.64 8.83 5.25 0.66 139.14

R_701_01 Nordkappbukta 16.06 11.09 10.03 0.83 503.93

R_701_02 Nordkappbukta 12.14 11.00 5.74 0.63 216.23

R_701_03 Nordkappbukta 16.90 11.24 8.30 0.71 444.76

R_701_04 Nordkappbukta 13.55 9.84 6.91 0.71 259.90

R_701_05 Nordkappbukta 12.00 8.21 5.40 0.67 150.08

R_701_06 Nordkappbukta 9.21 9.03 5.70 0.73 133.73

R_701_07 Nordkappbukta 22.20 15.57 5.85 0.46 570.42

R_701_08 Nordkappbukta 11.11 7.13 4.85 0.67 108.38

R_701_09 Nordkappbukta 19.30 17.30 6.82 0.52 642.37

R_701_10 Nordkappbukta 13.37 7.64 3.50 0.49 100.85

R_701_11 Nordkappbukta 12.36 9.03 5.34 0.63 168.13

R_701_12 Nordkappbukta 12.32 9.79 3.82 0.49 129.97

R_701_13 Nordkappbukta 11.07 9.49 4.34 0.56 128.62

R_701_14 Nordkappbukta 17.90 9.75 6.72 0.64 330.84

R_701_15 Nordkappbukta 11.13 10.16 5.54 0.65 176.72

R_701_16 Nordkappbukta 12.75 8.99 5.37 0.63 173.64

R_701_18 Nordkappbukta 9.21 7.25 2.46 0.45 46.34

R_701_19 Nordkappbukta 6.71 5.68 4.00 0.75 43.01

R_701_20 Nordkappbukta 9.00 4.56 4.32 0.77 50.01

R_701_21 Nordkappbukta 6.79 4.15 1.39 0.41 11.05

R_701_22 Nordkappbukta 5.96 2.90 2.68 0.75 13.07

R_701_23 Nordkappbukta 8.17 5.67 4.57 0.77 59.72

R_704_01 Nordkappbukta 3.97 2.88 1.42 0.56 4.58

R_704_02 Nordkappbukta 4.15 2.52 1.52 0.60 4.48

R_704_03 Nordkappbukta 5.59 3.53 1.29 0.44 7.18

R_704_04 Nordkappbukta 5.66 3.83 3.26 0.79 19.94

R_704_05 Nordkappbukta 4.85 4.32 2.46 0.66 14.54

R_704_06 Nordkappbukta 2.94 2.23 1.94 0.83 3.59

R_704_07 Nordkappbukta 2.71 2.25 1.48 0.71 2.55

R_704_08 Nordkappbukta 4.37 2.63 1.68 0.63 5.45

R_704_09 Nordkappbukta 4.13 3.44 3.41 0.94 13.67

R_704_10 Nordkappbukta 3.47 2.19 1.49 0.66 3.19

R_704_11 Nordkappbukta 6.06 3.72 2.25 0.61 14.31

R_704_12 Nordkappbukta 8.21 5.92 1.80 0.41 24.68

R_704_13 Nordkappbukta 6.67 3.81 1.41 0.43 10.11

R_704_14 Nordkappbukta 8.21 5.33 2.31 0.50 28.52

R_704_15 Nordkappbukta 5.11 3.79 1.93 0.58 10.54

R_704_16 Nordkappbukta 5.32 3.41 1.77 0.56 9.06

R_704_17 Nordkappbukta 5.00 3.26 2.52 0.73 11.59

R_704_18 Nordkappbukta 5.74 4.02 3.19 0.76 20.76

R_704_19 Nordkappbukta 5.67 4.99 1.47 0.42 11.73

R_704_20 Nordkappbukta 3.54 2.63 2.49 0.87 6.54 Appendix C

Rhodolith # Site Long axis (L) Intermediate axis (I) Short axis (S) Sphericity Size [cm] [cm] [cm] [(S2/LI)0.5] [(LIS/4π)0.5]

R_704_21 Nordkappbukta 4.96 4.64 3.54 0.82 22.98

R_711_01 Nordkappbukta 10.46 7.89 7.24 0.86 168.56

R_711_02 Nordkappbukta 10.46 7.85 5.70 0.73 132.03

R_711_03 Nordkappbukta 13.75 9.34 6.31 0.68 228.60

R_711_04 Nordkappbukta 10.30 10.24 5.00 0.62 148.77

R_711_05 Nordkappbukta 10.28 7.55 5.41 0.72 118.45

R_711_06 Nordkappbukta 11.64 7.50 6.02 0.75 148.25

R_711_07 Nordkappbukta 16.20 10.69 5.66 0.57 276.51

R_711_08 Nordkappbukta 11.46 9.53 8.71 0.89 268.34

R_711_09 Nordkappbukta 9.62 8.16 5.36 0.72 118.69

R_711_10 Nordkappbukta 10.63 8.68 5.10 0.66 132.75

R_711_11 Nordkappbukta 9.10 7.85 5.43 0.74 109.42

R_711_12 Nordkappbukta 14.21 11.47 6.24 0.62 286.90

R_711_13 Nordkappbukta 9.83 6.93 4.02 0.62 77.25

R_711_14 Nordkappbukta 10.36 6.18 4.77 0.71 86.15

R_711_15 Nordkappbukta 9.03 7.96 3.05 0.51 61.84

R_711_16 Nordkappbukta 11.91 10.73 7.60 0.77 273.98

R_711_17 Nordkappbukta 9.31 8.15 4.30 0.62 92.04

R_711_18 Nordkappbukta 9.56 7.07 6.04 0.81 115.16

R_711_19 Nordkappbukta 12.59 5.28 5.16 0.74 96.76

R_711_20 Nordkappbukta 9.96 6.77 3.43 0.56 65.24

R_711_21 Nordkappbukta 10.63 8.20 7.24 0.84 178.02

R_711_22 Nordkappbukta 11.97 9.87 9.05 0.89 301.62

R_711_23 Nordkappbukta 9.64 7.71 5.76 0.76 120.77

R_711_24 Nordkappbukta 15.20 14.50 9.29 0.73 577.59

R_711_25 Nordkappbukta 9.81 9.32 9.17 0.97 236.51

R_711_26 Nordkappbukta 8.18 6.36 4.50 0.73 66.04

R_711_27 Nordkappbukta 5.86 4.05 4.02 0.88 26.91

R_711_28 Nordkappbukta 6.30 5.33 4.60 0.86 43.57

R_711_29 Nordkappbukta 5.29 4.84 3.25 0.74 23.47

R_711_30 Nordkappbukta 6.05 5.03 4.41 0.86 37.86

R_711_31 Nordkappbukta 10.85 9.00 1.64 0.30 45.18

R_711_32 Nordkappbukta 8.27 6.52 5.34 0.81 81.22

R_711_33 Nordkappbukta 10.01 7.33 4.27 0.63 88.38

R_711_34 Nordkappbukta 5.41 4.41 2.69 0.67 18.10

R_711_35 Nordkappbukta 7.93 4.33 4.59 0.85 44.46

R_711_36 Nordkappbukta 8.13 6.84 4.84 0.75 75.93

R_711_37 Nordkappbukta 10.14 7.05 2.87 0.49 57.88

R_711_38 Nordkappbukta 7.16 6.01 4.24 0.75 51.47

R_711_39 Nordkappbukta 4.33 3.13 3.30 0.93 12.62

R_711_40 Nordkappbukta 7.09 4.90 3.77 0.74 36.95

R_711_41 Nordkappbukta 6.57 3.50 3.28 0.78 21.28

R_711_42 Nordkappbukta 8.04 6.88 4.74 0.74 73.96

R_711_43 Nordkappbukta 12.80 11.76 9.06 0.82 384.72

R_714_01 Nordkappbukta 9.11 7.83 6.40 0.83 128.78

R_714_02 Nordkappbukta 9.83 9.67 5.00 0.64 134.07

R_714_03 Nordkappbukta 10.69 8.22 6.24 0.76 154.68

R_714_04 Nordkappbukta 10.09 7.32 7.08 0.88 147.51

R_714_05 Nordkappbukta 7.57 6.85 6.32 0.92 92.45

R_714_06 Nordkappbukta 7.42 6.73 5.29 0.82 74.52

R_714_07 Nordkappbukta 8.05 7.39 6.26 0.87 105.05

R_714_08 Nordkappbukta 7.17 6.35 5.21 0.84 66.92

R_714_09 Nordkappbukta 7.47 6.87 5.83 0.87 84.40 Appendix C

Rhodolith # Site Long axis (L) Intermediate axis (I) Short axis (S) Sphericity Size [cm] [cm] [cm] [(S2/LI)0.5] [(LIS/4π)0.5]

R_714_10 Nordkappbukta 7.39 7.49 5.59 0.83 87.28

R_714_11 Nordkappbukta 7.95 7.17 5.73 0.83 92.14

R_714_12 Nordkappbukta 8.44 5.79 6.13 0.92 84.50

R_714_13 Nordkappbukta 7.50 6.98 5.61 0.84 82.85

R_714_14 Nordkappbukta 7.88 6.83 5.83 0.86 88.51

R_714_15 Nordkappbukta 9.29 7.50 4.90 0.70 96.31

R_714_16 Nordkappbukta 7.42 6.08 6.05 0.93 76.99

R_714_17 Nordkappbukta 9.46 7.38 7.25 0.91 142.78

R_714_18 Nordkappbukta 8.35 6.45 5.88 0.86 89.33

R_714_19 Nordkappbukta 8.94 7.64 6.50 0.85 125.24

R_714_20 Nordkappbukta 8.75 7.54 5.94 0.81 110.55

R_714_21 Nordkappbukta 9.00 7.05 6.50 0.87 116.34

R_714_22 Nordkappbukta 8.21 7.21 4.53 0.70 75.64

R_714_23 Nordkappbukta 7.50 7.58 5.78 0.84 92.69

R_714_24 Nordkappbukta 9.36 6.96 6.65 0.88 122.21

R_714_25 Nordkappbukta 7.78 6.72 5.26 0.81 77.58

R_714_26 Nordkappbukta 7.88 6.04 4.86 0.79 65.25

R_714_27 Nordkappbukta 6.40 5.30 5.27 0.94 50.43

R_714_28 Nordkappbukta 6.56 5.97 4.53 0.81 50.05

R_714_29 Nordkappbukta 7.10 6.35 5.30 0.85 67.41

R_714_30 Nordkappbukta 5.97 4.67 4.16 0.85 32.72

R_714_31 Nordkappbukta 6.97 4.99 4.28 0.81 41.99

R_714_32 Nordkappbukta 6.77 5.60 4.90 0.86 52.40

R_714_33 Nordkappbukta 5.70 5.28 4.11 0.82 34.89

R_714_34 Nordkappbukta 5.29 4.93 4.49 0.92 33.03

R_714_35 Nordkappbukta 7.25 6.73 4.89 0.79 67.31

R_714_36 Nordkappbukta 7.78 6.31 4.87 0.78 67.44

R_714_37 Nordkappbukta 7.30 6.29 4.67 0.78 60.49

R_714_38 Nordkappbukta 5.40 5.07 4.15 0.86 32.05

R_714_39 Nordkappbukta 6.50 6.30 5.00 0.85 57.76

R_714_40 Nordkappbukta 8.55 7.34 5.00 0.74 88.52

R_714_41 Nordkappbukta 7.10 5.93 4.43 0.78 52.62

R_714_42 Nordkappbukta 6.18 4.46 4.03 0.84 31.33

R_757_01 Isfjorden 15.44 9.71 2.19 0.32 92.62

R_757_02 Isfjorden 19.00 11.63 5.03 0.49 313.54

R_757_03 Isfjorden 12.02 10.12 7.82 0.80 268.34

R_757_04 Isfjorden 10.22 7.95 3.94 0.58 90.30

R_757_05 Isfjorden 8.64 5.34 4.91 0.81 63.90

R_757_06 Isfjorden 13.18 10.70 4.19 0.50 166.69

R_757_07 Isfjorden 13.00 8.72 5.35 0.63 171.08

R_757_08 Isfjorden 9.96 8.84 3.49 0.52 86.68

R_757_09 Isfjorden 9.13 7.08 5.85 0.81 106.67

R_757_10 Isfjorden 17.50 11.92 7.40 0.64 435.45

R_757_11 Isfjorden 12.66 11.99 4.99 0.55 213.67

R_757_12 Isfjorden 14.48 9.96 3.72 0.46 151.34

R_757_13 Isfjorden 14.10 10.46 5.44 0.59 226.33

R_757_14 Isfjorden 10.49 9.14 4.14 0.56 111.97

R_757_15 Isfjorden 12.54 10.25 4.02 0.50 145.76

R_757_16 Isfjorden 15.09 8.95 2.45 0.35 93.34

R_757_17 Isfjorden 11.78 7.30 5.95 0.74 144.34

R_757_18 Isfjorden 12.58 9.63 5.31 0.62 181.47

R_757_19 Isfjorden 13.21 8.69 4.70 0.58 152.20

R_757_20 Isfjorden 12.85 9.83 3.73 0.48 132.91 Appendix C

Rhodolith # Site Long axis (L) Intermediate axis (I) Short axis (S) Sphericity Size [cm] [cm] [cm] [(S2/LI)0.5] [(LIS/4π)0.5]

R_757_21 Isfjorden 10.61 7.89 4.03 0.58 95.17

R_757_22 Isfjorden 11.87 9.20 3.70 0.50 113.98

R_757_23 Isfjorden 9.29 6.06 5.81 0.84 92.27

R_757_24 Isfjorden 5.94 5.51 2.16 0.52 19.94

R_757_25 Isfjorden 10.00 6.10 3.09 0.54 53.17

R_757_26 Isfjorden 7.90 6.10 4.40 0.74 59.81

R_757_27 Isfjorden 6.12 3.90 2.90 0.71 19.53

R_757_28 Isfjorden 8.86 6.87 2.62 0.48 44.99

R_757_29 Isfjorden 10.03 5.75 5.05 0.76 82.16

R_757_30 Isfjorden 7.15 5.77 3.65 0.69 42.48

R_757_31 Isfjorden 6.87 5.34 4.46 0.82 46.16

R_757_32 Isfjorden 7.13 5.92 4.84 0.82 57.63

R_757_33 Isfjorden 6.38 5.05 3.50 0.72 31.81

R_757_34 Isfjorden 7.73 5.05 3.81 0.72 41.96

R_757_35 Isfjorden 7.94 3.80 1.21 0.36 10.30

R_757_36 Isfjorden 6.73 4.54 4.14 0.82 35.68

R_757_37 Isfjorden 8.38 4.66 4.03 0.75 44.39 Appendix D

Appendix D

Raw data rhodolith surface- and protuberance-measurements

Rhodolith # Site depth [m] protuberances surface protuberances/surface protuberances/surface [n] [cm2] [n cm-2] [n m-2]

R_637_03 Krossfjorden 50 69 14.6 4.73 47267

R_637_05 Krossfjorden 50 75 11.5 6.53 65266

R_637_11 Krossfjorden 50 96 12.2 7.87 78728

R_637_12 Krossfjorden 50 63 12.6 5.01 50112

R_637_16 Krossfjorden 50 79 15.9 4.97 49665

R_637_25 Krossfjorden 50 63 9.6 6.53 65335

R_637_28 Krossfjorden 50 41 8.4 4.88 48813

R_637_29 Krossfjorden 50 64 9.9 6.43 64341

R_637_31 Krossfjorden 50 80 9.5 8.40 84043

R_637_35 Krossfjorden 50 53 9.3 5.71 57106

R_637_36 Krossfjorden 50 45 6.7 6.72 67223

R_637_37 Krossfjorden 50 64 9.2 6.96 69636

R_637_38 Krossfjorden 50 26 7.3 3.54 35375

R_637_43 Krossfjorden 50 72 7.2 9.95 99539

R_637_44 Krossfjorden 50 26 8.5 3.04 30446

R_637_49 Krossfjorden 50 60 7.3 8.20 82023

R_637_51 Krossfjorden 50 44 6.4 6.86 68560

R_637_55 Krossfjorden 50 51 5.8 8.77 87678

R_637_57 Krossfjorden 50 36 6.1 5.92 59229

R_637_58 Krossfjorden 50 31 6.0 5.21 52071

R_637_61 Krossfjorden 50 46 6.5 7.11 71096

R_637_62 Krossfjorden 50 82 5.8 14.05 140485

R_637_65 Krossfjorden 50 29 4.0 7.27 72670

R_637_71 Krossfjorden 50 57 5.2 10.97 109716

R_637_79 Krossfjorden 50 52 4.7 11.08 110809

R_637_80 Krossfjorden 50 38 4.0 9.46 94612

R_637_83 Krossfjorden 50 33 4.4 7.42 74219

R_637_85 Krossfjorden 50 38 3.4 11.14 111441

R_637_88 Krossfjorden 50 53 3.7 14.18 141810

R_637_92 Krossfjorden 50 30 4.2 7.12 71189

R_637_96 Krossfjorden 50 45 5.2 8.58 85821

R_644_01 Krossfjorden 41 116 20.5 5.66 56562

R_644_02 Krossfjorden 41 32 11.6 2.76 27573

R_644_03 Krossfjorden 41 118 11.2 10.52 105201

R_644_04 Krossfjorden 41 108 16.1 6.71 67051

R_644_05 Krossfjorden 41 147 24.1 6.10 60966

R_644_06 Krossfjorden 41 118 19.8 5.97 59673

R_644_07 Krossfjorden 41 66 13.0 5.07 50689

R_644_08 Krossfjorden 41 144 17.8 8.10 80983

R_644_09 Krossfjorden 41 26 8.7 2.99 29859

R_644_10 Krossfjorden 41 116 26.2 4.43 44341

R_644_11 Krossfjorden 41 95 14.9 6.38 63798

R_644_12 Krossfjorden 41 118 18.3 6.46 64615

R_644_13 Krossfjorden 41 70 13.8 5.07 50714

R_644_14 Krossfjorden 41 143 20.7 6.91 69108 Appendix D

Rhodolith # Site depth [m] protuberances surface protuberances/surface protuberances/surface [n] [cm2] [n cm-2] [n m-2]

R_644_15 Krossfjorden 41 102 44.5 2.29 22903

R_652_01 Krossfjorden 47 167 32.6 5.12 51241

R_652_02 Krossfjorden 47 100 16.8 5.94 59363

R_684_01 Mosselbukta 44 172 59.9 2.87 28737

R_684_04 Mosselbukta 44 215 51.9 4.14 41436

R_684_07 Mosselbukta 44 270 94.7 2.85 28499

R_684_14 Mosselbukta 44 141 62.4 2.26 22580

R_684_17 Mosselbukta 44 200 99.0 2.02 20195

R_684_18 Mosselbukta 44 169 42.5 3.98 39777

R_684_19 Mosselbukta 44 128 47.6 2.69 26915

R_684_24 Mosselbukta 44 160 74.0 2.16 21630

R_684_27 Mosselbukta 44 105 36.8 2.85 28526

R_684_29 Mosselbukta 44 224 54.5 4.11 41120

R_684_33 Mosselbukta 44 206 34.3 6.01 60094

R_684_35 Mosselbukta 44 101 39.4 2.56 25619

R_684_40 Mosselbukta 44 261 106.6 2.45 24495

R_684_49 Mosselbukta 44 219 49.8 4.39 43946

R_684_52 Mosselbukta 44 257 92.1 2.79 27898

R_684_53 Mosselbukta 44 149 41.0 3.64 36359

R_684_54 Mosselbukta 44 128 46.5 2.75 27501

R_684_55 Mosselbukta 44 144 48.2 2.99 29899

R_684_56 Mosselbukta 44 112 57.8 1.94 19380

R_684_63 Mosselbukta 44 160 75.0 2.13 21326

R_684_65 Mosselbukta 44 199 63.5 3.13 31338

R_684_66 Mosselbukta 44 263 52.2 5.04 50374

R_684_69 Mosselbukta 44 319 124.6 2.56 25610

R_684_72 Mosselbukta 44 136 40.4 3.37 33676

R_684_74 Mosselbukta 44 194 62.1 3.13 31259

R_684_75 Mosselbukta 44 202 84.9 2.38 23790

R_684_76 Mosselbukta 44 167 59.0 2.83 28308

R_701_03 Nordkappbukta 38 211 176.3 1.20 11967

R_701_06 Nordkappbukta 38 151 62.5 2.42 24172

R_701_10 Nordkappbukta 38 170 83.3 2.04 20409

R_701_12 Nordkappbukta 38 197 73.1 2.69 26939

R_701_13 Nordkappbukta 38 316 77.3 4.09 40903

R_701_15 Nordkappbukta 38 205 113.1 1.81 18133

R_701_16 Nordkappbukta 38 131 86.7 1.51 15117

R_701_19 Nordkappbukta 38 63 24.6 2.56 25581

R_701_20 Nordkappbukta 38 80 50.0 1.60 16001

R_701_23 Nordkappbukta 38 120 50.1 2.39 23950

R_711_01 Nordkappbukta 45 192 77.8 2.47 24670

R_711_02 Nordkappbukta 45 159 71.2 2.23 22341

R_711_05 Nordkappbukta 45 107 59.5 1.80 17981

R_711_06 Nordkappbukta 45 109 76.0 1.44 14351

R_711_08 Nordkappbukta 45 240 91.2 2.63 26327

R_711_10 Nordkappbukta 45 123 60.3 2.04 20401

R_711_11 Nordkappbukta 45 153 65.8 2.33 23270

R_711_13 Nordkappbukta 45 45 50.3 0.89 8946

R_711_15 Nordkappbukta 45 201 58.5 3.44 34370

R_711_16 Nordkappbukta 45 196 101.7 1.93 19278

R_711_18 Nordkappbukta 45 173 54.1 3.20 31951

R_711_19 Nordkappbukta 45 128 61.8 2.07 20717

R_711_20 Nordkappbukta 45 148 45.4 3.26 32593 Appendix D

Rhodolith # Site depth [m] protuberances surface protuberances/surface protuberances/surface [n] [cm2] [n cm-2] [n m-2]

R_711_22 Nordkappbukta 45 208 103.5 2.01 20089

R_711_23 Nordkappbukta 45 105 53.5 1.96 19624

R_711_25 Nordkappbukta 45 192 83.0 2.31 23123

R_711_26 Nordkappbukta 45 61 35.8 1.71 17059

R_711_27 Nordkappbukta 45 48 22.4 2.15 21471

R_711_29 Nordkappbukta 45 55 18.7 2.94 29449

R_711_31 Nordkappbukta 45 118 72.5 1.63 16277

R_711_32 Nordkappbukta 45 46 43.5 1.06 10585

R_711_34 Nordkappbukta 45 76 17.4 4.37 43686

R_711_35 Nordkappbukta 45 132 28.2 4.69 46883

R_711_36 Nordkappbukta 45 138 44.9 3.07 30721

R_711_37 Nordkappbukta 45 66 52.2 1.26 12644

R_711_41 Nordkappbukta 45 55 25.8 2.13 21336

R_711_42 Nordkappbukta 45 76 39.6 1.92 19199

R_711_43 Nordkappbukta 45 403 113.1 3.56 35629

R_714_01 Nordkappbukta 27 135 61.7 2.19 21884

R_714_02 Nordkappbukta 27 97 70.1 1.38 13838

R_714_03 Nordkappbukta 27 144 75.7 1.90 19025

R_714_04 Nordkappbukta 27 75 56.9 1.32 13170

R_714_05 Nordkappbukta 27 138 44.5 3.10 31010

R_714_06 Nordkappbukta 27 108 43.9 2.46 24626

R_714_07 Nordkappbukta 27 137 49.1 2.79 27900

R_714_09 Nordkappbukta 27 113 39.8 2.84 28424

R_714_10 Nordkappbukta 27 158 50.7 3.12 31154

R_714_13 Nordkappbukta 27 96 45.1 2.13 21268

R_714_14 Nordkappbukta 27 135 42.4 3.18 31839

R_714_15 Nordkappbukta 27 124 57.1 2.17 21703

R_714_16 Nordkappbukta 27 44 44.2 1.00 9953

R_714_17 Nordkappbukta 27 76 69.3 1.10 10965

R_714_18 Nordkappbukta 27 89 46.4 1.92 19161

R_714_19 Nordkappbukta 27 144 61.2 2.35 23512

R_714_20 Nordkappbukta 27 84 62.6 1.34 13417

R_714_21 Nordkappbukta 27 106 55.5 1.91 19085

R_714_22 Nordkappbukta 27 86 47.3 1.82 18182

R_714_23 Nordkappbukta 27 231 53.1 4.35 43489

R_714_24 Nordkappbukta 27 264 56.2 4.70 46987

R_714_25 Nordkappbukta 27 186 51.8 3.59 35909

R_714_26 Nordkappbukta 27 126 41.0 3.07 30740

R_714_27 Nordkappbukta 27 129 30.0 4.30 43024

R_714_28 Nordkappbukta 27 148 29.4 5.04 50375

R_714_29 Nordkappbukta 27 204 41.0 4.97 49728

R_714_30 Nordkappbukta 27 88 22.4 3.93 39257

R_714_31 Nordkappbukta 27 170 33.0 5.15 51482

R_714_32 Nordkappbukta 27 143 37.5 3.81 38092

R_714_33 Nordkappbukta 27 157 24.1 6.51 65088

R_714_34 Nordkappbukta 27 102 19.9 5.13 51339

R_714_36 Nordkappbukta 27 133 40.2 3.31 33070

R_714_37 Nordkappbukta 27 97 33.1 2.93 29332

R_714_38 Nordkappbukta 27 135 22.8 5.93 59313

R_714_39 Nordkappbukta 27 112 33.6 3.33 33331

R_714_41 Nordkappbukta 27 103 36.3 2.84 28389

R_714_42 Nordkappbukta 27 121 25.3 4.78 47795

R_757_04 Floskjeret 45 239 76.8 3.11 31134 Appendix D

Rhodolith # Site depth [m] protuberances surface protuberances/surface protuberances/surface [n] [cm2] [n cm-2] [n m-2]

R_757_05 Floskjeret 45 240 57.0 4.21 42116

R_757_06 Floskjeret 45 332 94.2 3.52 35239

R_757_07 Floskjeret 45 419 109.8 3.82 38163

R_757_16 Floskjeret 45 313 140.6 2.23 22267

R_757_17 Floskjeret 45 243 70.8 3.43 34341

R_757_23 Floskjeret 45 117 48.6 2.41 24082

R_757_25 Floskjeret 45 66 45.7 1.45 14454

R_757_26 Floskjeret 45 95 44.0 2.16 21566

R_757_29 Floskjeret 45 138 50.5 2.73 27347

R_757_30 Floskjeret 45 64 35.0 1.83 18294

R_757_31 Floskjeret 45 144 37.4 3.85 38504

R_757_32 Floskjeret 45 171 40.2 4.26 42576

R_757_33 Floskjeret 45 118 21.2 5.56 55636

R_757_34 Floskjeret 45 124 29.3 4.23 42338

R_757_36 Floskjeret 45 122 27.2 4.48 44811

R_757_37 Floskjeret 45 181 38.1 4.75 47462 Appendix E

Appendix E

Raw data protuberance weight- and increment-measurements

Protuberance # Weight Lenght Bands Weight/band Lenght/band [mg] [mm] [n] [mg n-1] [µm n-1]

P_637_3_1 52.3 5.4 14 3.7 382

P_637_3_2 65.3 5.5 20 3.3 274

P_637_3_3 80 5.6 23 3.5 245

P_637_3_4 88.3 6.6 27 3.3 244

P_637_3_5 63.9 5.4 20 3.2 269

P_637_5_1 17.5 2.4 7 2.5 344

P_637_5_2 13.3 2.5 7 1.9 357

P_637_5_3 18 2.7 7 2.6 383

P_637_5_4 21.4 2.8 8 2.7 345

P_637_5_5 25.9 3.1 7 3.7 447

P_637_16_1 113.8 5.8 27 4.2 214

P_637_16_2 73.3 5.7 21 3.5 270

P_637_16_3 64 5.1 23 2.8 223

P_637_16_4 34.7 3.7 15 2.3 249

P_637_16_5 35.2 3.8 19 1.9 197

P_637_51_1 57.6 4.7 23 2.5 203

P_637_51_2 55.1 4.6 21 2.6 220

P_637_51_3 69.8 5.1 23 3.0 221

P_637_51_4 67.7 5.3 19 3.6 281

P_637_51_5 88.4 6.1 25 3.5 245

P_637_71_1 21.8 3.0 12 1.8 249

P_637_71_2 23.8 3.4 11 2.2 305

P_637_71_3 30.1 3.5 13 2.3 268

P_637_71_4 23.9 2.8 11 2.2 255

P_637_71_5 42.7 4.3 15 2.8 283

P_637_79_1 14.3 2.9 10 1.4 291

P_637_79_2 13.2 2.4 9 1.5 264

P_637_79_3 15.1 3.0 9 1.7 338

P_637_79_4 12.1 2.3 7 1.7 334

P_637_79_5 21.9 3.5 10 2.2 352

P_637_96_1 12.3 2.3 7 1.8 326

P_637_96_2 13.9 2.5 6 2.3 420

P_637_96_3 18.1 2.8 9 2.0 308

P_637_96_4 15.6 2.6 10 1.6 263

P_637_96_5 18.6 2.9 11 1.7 259

P_644_1_1 71.4 5.1 24 3.0 214

P_644_1_2 93.1 5.8 27 3.4 214

P_644_1_3 31.6 3.6 17 1.9 211

P_644_1_4 100.5 4.7 24 4.2 196

P_644_1_5 69.4 5.2 23 3.0 224

P_644_3_1 61.6 4.6 23 2.7 200

P_644_3_2 84.9 4.5 25 3.4 181

P_644_3_3 49.5 4.4 22 2.3 199

P_644_3_4 55.9 4.3 23 2.4 189

P_644_3_5 72.9 5.0 23 3.2 217 Appendix E

Protuberance # Weight Lenght Bands Weight/band Lenght/band [mg] [mm] [n] [mg n-1] [µm n-1]

P_644_4_1 49.9 4.2 19 2.6 220

P_644_4_2 64.3 4.9 21 3.1 233

P_644_4_3 63.3 6.0 21 3.0 287

P_644_4_4 81.9 5.6 24 3.4 232

P_644_4_5 50.2 4.9 24 2.1 202

P_644_5_1 100.6 6.9 31 3.2 221

P_644_5_2 84 5.5 27 3.1 202

P_644_5_3 82.5 6.0 28 2.9 213

P_644_5_4 66.6 4.8 26 2.6 183

P_644_5_5 71.5 5.1 26 2.8 197

P_644_6_1 89 5.2 21 4.2 245

P_644_6_2 81.1 4.9 22 3.7 223

P_644_6_3 32.7 3.7 18 1.8 203

P_644_6_4 61.8 4.9 24 2.6 205

P_644_6_5 51.4 5.5 18 2.9 307

P_644_7_1 29.7 3.2 14 2.1 230

P_644_7_2 46.4 4.4 22 2.1 200

P_644_7_3 50.3 4.9 19 2.6 260

P_644_7_4 38.8 4.2 18 2.2 233

P_644_7_5 41.2 4.0 15 2.7 269

P_652_1_1 43.7 4.1 18 2.4 229

P_652_1_2 161.2 7.6 41 3.9 186

P_652_1_3 50.3 3.8 17 3.0 221

P_652_1_4 94.2 6.9 29 3.2 239

P_652_1_5 68 5.4 27 2.5 198

P_652_2_1 101.8 8.1 32 3.2 252

P_652_2_2 119.1 6.9 36 3.3 192

P_652_2_3 122.5 7.1 32 3.8 220

P_652_2_4 115.8 6.5 38 3.0 172

P_652_2_5 58.4 4.8 26 2.2 183

P_684_9_1 59.1 3.9 18 3.3 214

P_684_9_2 46 4.0 20 2.3 202

P_684_9_3 75.3 4.5 19 4.0 236

P_684_9_4 94.3 4.7 26 3.6 182

P_684_9_5 56.2 4.8 17 3.3 282

P_684_18_1 65.7 5.4 23 2.9 233

P_684_18_2 136.5 6.5 28 4.9 230

P_684_18_3 135.4 5.9 27 5.0 219

P_684_18_4 127.5 6.3 25 5.1 252

P_684_18_5 69.4 5.1 21 3.3 243

P_684_29_1 94.2 5.6 24 3.9 233

P_684_29_2 123.2 7.3 31 4.0 235

P_684_29_3 244.3 5.8 41 6.0 142

P_684_29_4 61 4.9 23 2.7 211

P_684_29_5 212.3 6.3 34 6.2 184

P_684_35_1 65.4 5.6 27 2.4 207

P_684_35_2 51.5 6.3 18 2.9 351

P_684_35_3 202.5 8.5 32 6.3 267

P_684_35_4 94 7.6 27 3.5 280

P_684_35_5 135.6 6.5 26 5.2 251

P_684_36_1 99.6 5.1 26 3.8 198

P_684_36_2 65.8 4.6 22 3.0 207

P_684_36_3 231.1 5.4 32 7.2 170 Appendix E

Protuberance # Weight Lenght Bands Weight/band Lenght/band [mg] [mm] [n] [mg n-1] [µm n-1]

P_684_36_4 197.7 7.7 30 6.6 256

P_684_36_5 86.4 6.4 24 3.6 266

P_684_45_1 329 10.3 53 6.2 195

P_684_45_2 573 11.7 73 7.8 160

P_684_45_3 170.2 5.1 33 5.2 155

P_684_45_4 336.5 7.7 40 8.4 191

P_684_45_5 267.1 6.9 38 7.0 183

P_684_46_1 211.7 6.5 32 6.6 204

P_684_46_2 87.5 4.5 25 3.5 178

P_684_46_3 100.1 4.6 29 3.5 159

P_684_46_4 81.7 4.8 33 2.5 144

P_684_46_5 59.5 3.9 26 2.3 152

P_684_51_1 111.9 4.3 25 4.5 172

P_684_51_2 77.2 6.4 32 2.4 200

P_684_51_3 141.8 5.4 24 5.9 226

P_684_51_4 54.3 5.0 25 2.2 199

P_684_51_5 52.2 4.0 19 2.7 210

P_684_65_1 167.5 7.0 28 6.0 248

P_684_65_2 72.4 6.0 26 2.8 232

P_684_65_3 119 4.5 26 4.6 174

P_684_65_4 107.3 4.8 27 4.0 178

P_684_65_5 236.5 8.2 43 5.5 190

P_684_72_1 72.5 4.8 25 2.9 191

P_684_72_2 130.8 6.0 30 4.4 200

P_684_72_3 97.5 5.3 26 3.8 203

P_684_72_4 68 5.6 23 3.0 243

P_684_72_5 49.7 3.7 21 2.4 177

P_684_74_1 129.6 7.0 32 4.1 219

P_684_74_2 71.8 6.9 29 2.5 236

P_684_74_3 69.1 5.4 25 2.8 214

P_684_74_4 202.3 10.2 41 4.9 249

P_684_74_5 150.5 6.3 29 5.2 217

P_684_75_1 119.9 8.3 36 3.3 231

P_684_75_2 175.5 11.6 38 4.6 304

P_684_75_3 93 7.4 29 3.2 256

P_684_75_4 57.9 6.4 19 3.0 336

P_684_75_5 131.8 9.8 35 3.8 279

P_701_6_1 173.4 11.4 47 3.7 243

P_701_6_2 186.4 9.0 50 3.7 181

P_701_6_3 303.5 10.5 60 5.1 175

P_701_6_4 788.2 19.5 92 8.6 212

P_701_6_5 640.4 18.6 76 8.4 245

P_701_7_1 160.7 7.5 43 3.7 175

P_701_7_2 154.7 5.8 26 6.0 223

P_701_7_3 99.3 6.3 35 2.8 179

P_701_7_4 252.8 7.7 39 6.5 198

P_701_7_5 505 18.7 76 6.6 246

P_701_10_1 209.6 8.4 38 5.5 222

P_701_10_2 95.9 6.6 43 2.2 154

P_701_10_3 137.2 6.6 34 4.0 194

P_701_10_4 171.6 6.8 38 4.5 180

P_701_10_5 168 7.1 33 5.1 215

P_701_12_1 80.5 6.0 35 2.3 172 Appendix E

Protuberance # Weight Lenght Bands Weight/band Lenght/band [mg] [mm] [n] [mg n-1] [µm n-1]

P_701_12_2 480.6 13.0 64 7.5 203

P_701_12_3 102.2 8.0 39 2.6 206

P_701_12_4 114.6 5.0 30 3.8 166

P_701_12_5 156 6.4 36 4.3 176

P_701_13_1 74.5 5.5 27 2.8 202

P_701_13_2 169.4 6.7 36 4.7 185

P_701_13_3 34.2 3.9 22 1.6 176

P_701_13_4 285.5 8.8 52 5.5 170

P_701_13_5 61.1 5.6 26 2.4 214

P_701_16_1 62.7 5.6 18 3.5 311

P_701_16_2 152.4 8.5 35 4.4 243

P_701_16_3 106.4 4.6 27 3.9 171

P_701_16_4 196.6 4.9 31 6.3 158

P_701_16_5 112.8 4.4 18 6.3 242

P_701_23_1 418.6 10.3 56 7.5 185

P_701_23_2 113.5 6.6 38 3.0 174

P_701_23_3 77.5 6.1 39 2.0 157

P_701_23_4 176.8 8.9 34 5.2 262

P_701_23_5 147.6 7.4 46 3.2 160

P_711_6_1 799.1 14.4 96 8.3 150

P_711_6_2 237.1 9.8 57 4.2 172

P_711_6_3 104.6 5.7 34 3.1 168

P_711_6_4 385 9.6 55 7.0 174

P_711_6_5 88.8 4.8 35 2.5 137

P_711_11_1 172.7 7.9 43 4.0 183

P_711_11_2 287.4 9.7 44 6.5 219

P_711_11_3 200.1 7.1 50 4.0 142

P_711_11_4 137 7.6 48 2.9 158

P_711_11_5 235.3 9.4 62 3.8 152

P_711_23_1 147.1 6.5 32 4.6 203

P_711_23_2 342.8 10.7 52 6.6 206

P_711_23_3 305.5 9.8 58 5.3 169

P_711_23_4 129.3 6.9 40 3.2 174

P_711_23_5 168.2 7.5 36 4.7 207

P_711_25_1 175.5 6.8 34 5.2 200

P_711_25_2 199.6 8.9 52 3.8 171

P_711_25_3 370.3 12.7 57 6.5 223

P_711_25_4 222.5 9.8 37 6.0 264

P_711_25_5 461.1 13.7 69 6.7 198

P_711_36_1 159.5 6.9 45 3.5 152

P_711_36_2 201.3 7.4 48 4.2 155

P_711_36_3 176.9 8.4 47 3.8 179

P_711_36_4 393.4 10.9 69 5.7 158

P_711_36_5 144.5 7.7 44 3.3 174

P_711_40_1 251.7 11.8 61 4.1 193

P_711_40_2 504.9 12.0 74 6.8 162

P_711_40_3 204.5 7.7 43 4.8 179

P_711_40_4 366.5 10.8 71 5.2 151

P_711_40_5 244.3 9.4 51 4.8 184

P_711_43_1 260.5 10.1 52 5.0 195

P_711_43_2 249.2 10.2 50 5.0 204

P_711_43_3 181.3 9.2 40 4.5 230

P_711_43_4 340.6 9.7 51 6.7 191 Appendix E

Protuberance # Weight Lenght Bands Weight/band Lenght/band [mg] [mm] [n] [mg n-1] [µm n-1]

P_711_43_5 359.4 10.7 60 6.0 178

P_714_1_1 241 6.3 39 6.2 162

P_714_1_2 65.8 4.0 22 3.0 180

P_714_1_3 255.3 8.4 49 5.2 171

P_714_1_4 283.5 7.8 44 6.4 178

P_714_1_5 113 5.9 28 4.0 210

P_714_4_1 706.7 8.0 90 7.9 88

P_714_4_2 387.7 6.5 47 8.2 139

P_714_4_3 269.8 6.6 55 4.9 119

P_714_4_4 858.4 9.8 89 9.6 110

P_714_4_5 233.2 5.8 40 5.8 145

P_714_10_1 212.7 8.0 49 4.3 163

P_714_10_2 139.4 5.6 33 4.2 171

P_714_10_3 311.4 7.0 44 7.1 159

P_714_10_4 246.7 6.8 34 7.3 199

P_714_10_5 292.4 7.0 37 7.9 189

P_714_25_1 407.5 9.8 63 6.5 156

P_714_25_2 147.6 6.3 40 3.7 158

P_714_25_3 304.8 9.1 43 7.1 211

P_714_25_4 211.9 6.8 40 5.3 169

P_714_25_5 121.8 5.5 30 4.1 184

P_714_30_1 239.9 10.8 49 4.9 221

P_714_30_2 67.3 5.6 26 2.6 217

P_714_30_3 91.7 5.8 29 3.2 200

P_714_30_4 182.7 7.7 39 4.7 198

P_714_30_5 95.1 5.6 33 2.9 169

P_714_36_1 84.2 6.1 29 2.9 209

P_714_36_2 91.4 6.0 35 2.6 170

P_714_36_3 184.5 8.9 40 4.6 222

P_714_36_4 216.6 8.6 45 4.8 191

P_714_36_5 129.2 6.7 35 3.7 190

P_714_42_1 155.5 5.8 38 4.1 153

P_714_42_2 145.9 7.0 41 3.6 171

P_714_42_3 676.8 14.1 84 8.1 168

P_714_42_4 211.5 6.1 48 4.4 128

P_714_42_5 167.2 8.1 43 3.9 188

P_757_2_1 138 5.2 17 8.1 306

P_757_2_2 127 5.1 22 5.8 232

P_757_2_3 302 8.8 40 7.6 220

P_757_2_4 336 9.6 30 11.2 321

P_757_2_5 261 8.2 27 9.7 302

P_757_4_1 116 5.1 18 6.4 281

P_757_4_2 169 9.6 29 5.8 329

P_757_4_3 386 12.0 54 7.2 223

P_757_4_4 81 4.7 12 6.8 393

P_757_4_5 187 7.6 30 6.2 252

P_757_6_1 160 5.9 22 7.3 268

P_757_6_2 103 6.1 22 4.7 276

P_757_6_3 84 5.0 17 4.9 293

P_757_6_4 108 5.7 23 4.7 247

P_757_6_5 95 5.3 22 4.3 240

P_757_7_1 201 9.4 31 6.5 304

P_757_7_2 350 10.6 39 9.0 271 Appendix E

Protuberance # Weight Lenght Bands Weight/band Lenght/band [mg] [mm] [n] [mg n-1] [µm n-1]

P_757_7_3 51 4.4 17 3.0 258

P_757_7_4 194 9.0 28 6.9 323

P_757_7_5 87 6.5 22 4.0 297

P_757_16_1 652 11.3 64 10.2 177

P_757_16_2 90 4.2 20 4.5 211

P_757_16_3 145 6.6 32 4.5 208

P_757_16_4 268 8.6 35 7.7 247

P_757_16_5 228 8.0 36 6.3 223

P_757_17_1 91 5.1 22 4.1 230

P_757_17_2 85 4.7 18 4.7 260

P_757_17_3 132 6.3 24 5.5 264

P_757_17_4 187 6.1 26 7.2 234

P_757_17_5 153 6.1 28 5.5 218

P_757_26_1 91 5.1 25 3.6 204

P_757_26_2 58 5.0 18 3.2 275

P_757_26_3 115 6.4 16 7.2 402

P_757_26_4 118 5.8 24 4.9 243

P_757_26_5 87 6.0 23 3.8 262

P_757_31_1 178 7.1 33 5.4 216

P_757_31_2 227 9.1 34 6.7 269

P_757_31_3 127 5.7 21 6.0 273

P_757_31_4 77 5.1 19 4.1 268

P_757_31_5 159 6.7 29 5.5 230

P_757_32_1 117 6.1 21 5.6 290

P_757_32_2 85 5.3 18 4.7 297

P_757_32_3 119 6.1 19 6.3 323

P_757_32_4 128 5.8 20 6.4 291

P_757_32_5 103 5.8 21 4.9 276

P_757_33_1 93 6.0 19 4.9 314

P_757_33_2 51 5.0 16 3.2 313

P_757_33_3 43 4.2 14 3.1 301

P_757_33_4 100 6.5 17 5.9 385

P_757_33_5 94 6.3 24 3.9 261

P_757_34_1 63 5.4 21 3.0 255

P_757_34_2 60 4.9 20 3.0 246

P_757_34_3 115 6.5 19 6.1 340

P_757_34_4 116 6.5 23 5.0 283

P_757_34_5 98 5.7 21 4.7 272

P_757_36_1 207 9.1 32 6.5 284

P_757_36_2 204 8.5 35 5.8 243

P_757_36_3 162 7.5 23 7.1 328

P_757_36_4 138 7.6 26 5.3 290

P_757_36_5 96 5.7 18 5.3 318 Appendix F

Appendix F

Article citation concerning the Nordkappbukta rhodolith communities published in the ISI journal Phycologia

Appendix F comprises the article citation of “Rhodolith beds (Corallinales, Rhodophyta) and their physical and biological environment at 80°31’N in Nordkappbukta (Nordaustlandet, Svalbard Archipelago, Norway)” published in the ISI Journal Phycologia. The manuscript was submitted to Phycologia in July 2011 and accepted in October 2011. The attached article is to be cited as follows:

Teichert S., Woelkerling W., Rüggeberg A., Wisshak M., Piepenburg D., Meyerhöfer M., Form A., Büdenbender J. & Freiwald A. (2012) Rhodolith beds (Corallinales, Rhodo- phyta) and their physical and biological environment at 80°31’N in Nordkappbukta (Nord- austlandet, Svalbard Archipelago, Norway). Phycologia 51: 371-390.

The article is available from: http://www.phycologia.org DOI: http://dx.doi.org/10.2216/11-76.1