Description: Commercially exploited stocks that have experienced declines in population abundance have responded by altering life history traits of growth and maturation. Cod is not only becoming mature at an earlier age but also, the majority of the stock comprises fishes with no previous spawning experience. Actual fisheries management does not take in account qualitative differences within the spawning stock. If the stock responds to continued exploitation by shifting maturity to an earlier stage, fish will spawn at smaller sizes. They will produce smaller eggs, and consequently small and less viable larvae, so that the contribution to the spawning stock biomass will be less than expected. There are many advantages for delaying maturation: Larger and heavier fish will be better conditioned for spawning, have higher fecundity and larger eggs that are more viable. Harvesting at delayed recruitment enables the stock to maintain a larger SSB with an expanded age structure while supporting a sustainable fishery.

Description: Anthropogenic greenhouse gas emissions have been driving global climate change and they will continue to do so over the course of the 21st century. Most of the marine biosphere and especially coastal marine systems have suffered from high anthropogenic pressure per se and it is possible that the novel burden of very rapidly proceeding global climate change triggers shifts to alternative regimes and functioning in marine ecosystems. In the light of this background, my dissertation aims to contribute to the mechanistic understanding of global and local climate change effects on a common coastal marine seaweed (Fucus vesiculosus, Phaeophyceae) system of the Baltic Sea. The results of my experimental studies provide important mechanistic clues about the underlying direct and indirect effective pathways of environmental change in the studied seaweed system. To the best of my knowledge, it is one of the first studies which assess the seasonal variability of the same environmental factors on the same marine system over the course of one year. The detected context-dependency of global climate change effects within one ecosystem clearly shows that our understanding of the basic underlying ecosystem processes and patterns forms a prerequisite for testing, predicting and managing future ecological change in marine systems. Given that grazing forms a crucial ecological force in many coastal vegetated systems, the identified underlying mechanisms of change (top-down and bottom-up control) may allow reference to other similarly structured coastal systems. Importantly my findings point out, that ecological impacts of global climate change may be underestimated if local perturbation is disregarded and, thus, underline the chance and responsibility of local ecosystem management.

Description: The overall aim of my thesis was to improve our understanding of environmental drivers causing the dynamics of the Eastern Baltic cod reproduction habitat and to assess their relevance for a possible application in the sustainable stock management. A novel approach to map the reproductive habitat of Eastern Baltic cod, the Buoyancy Depending Reproductive Layers (BDRL), was developed and used to propose an alternative stock indicator, the effective Spawning Stock Biomass (eSSB). The eSSB was found to improve the fit of a recruitment model compared to the model using the conventional Spawning Stock Biomass. Oxygen depletion was also negatively impacting the available size of nursery areas for juveniles. The mechanism was able to partly explain the observed decline of the condition of juveniles in the nursery areas, because a high population density in the remaining habitat could increase the impact of density depending effects. Furthermore, by the application of the novel approach of BDRLs the spawning habitat was shown to be sensitive to eutrophication and that this sensitivity is strongly depending on the size of the female spawner using the habitat. As predicted, the BDRL approach was superior to the “classic” approach, the Reproductive Volume (RV) because it was more sensitive to environmental change, able to incorporate stock structure, was not overestimating the spawning habitat in the eastern spawning areas and could provide estimates of other stressors depending on the female spawner size. Due to a permanently installed measurement platform in the Arkona Basin, the new methodology could be used to establish a new environmental indicator on the spawning habitat conditions. It was recommended to be used in future stock assessments.

Description: The trace metal iron is considered to be the nutrient that limits marine primary production in one third of the global surface ocean (Martin, 1990; Boyd et al., 2007; Moore et al., 2013). It is also the nutrient that maintains future ocean fertility due to its irreplaceable role in the process of nitrogen fixation, which adds “new” nitrogen (another nutrient for phytoplankton) to the surface ocean (Raven, 1988; Kustka et al., 2003b; Zehr and Capone, 2020). Due to iron’s importance, it is not surprising that the demand for incorporating iron into global biogeochemical models is high. However, including iron in an earth system model has been shown to have no clear benefits with respect to model misfit against observational data (Nickelsen et al., 2015) . How smart is it then to introduce iron models into global biogeochemical models, when the benefits are not clearly identifiable? Especially, when the iron models perform poorly at reproducing observed iron patterns in the ocean (Tagliabue et al., 2016). The poor performance of iron models, coupled with their failure to improve biogeochemical tracer representation of the ocean, inspired this additional effort to identify the advantages of including iron in a global biogeochemical model, both for the preindustrial state and under conditions of a changing climate. The working hypothesis was that the relatively poor performance of iron models might come from inadequate model calibration. A first sensitivity study on biogeochemical model parameter values was conducted in order to identify key parameters for model calibration. It was found that while some of the parameters influence simulated nitrogen, phosphorus, and oxygen concentrations, few parameters influence simulated iron concentrations. This suggests that our modelling skill of the iron cycle is still limited and/or that the observational data base is insufficient for comprehensive model calibration so far. Thus it was decided not to include iron data in further model calibration. A model calibration framework (Kriest et al., 2017) was next applied to a hierarchy of global models with different implementations of iron; one without iron, one with prescribed iron concentrations, and another one with a dynamic iron cycle. Using calibration against global data sets of nitrogen, phosphorus, and oxygen, the misfit of each model was pushed to its minimum. It was found that under an assumed preindustrial steady state, the calibrated model with a full dynamic iron cycle has the lowest model misfit against observations (thus confirming the working hypothesis). It was also found that the calibrated model with a fully dynamic iron cycle has 50% less net primary production (which is closer to empirical estimations) compared to the calibrated model without iron. Finally, transient simulations for all calibrated models were integrated from their pre- industrial state until the end of the 21st century using an atmospheric CO2 concentration pathway consistent with a ’business-as-usual’ CO2 emission scenario. It was found that nitrogen fixation trends diverge among models. This divergence is caused by whether iron limits the productivity of the upwelling regions, e.g. in the eastern tropical Pacific. The export production in the eastern tropical Pacific (and other tropical upwelling regions) reacts differently to warming, depending on whether iron is a limiting nutrient. These different responses trigger a divergent chain of downstream responses that affect nitrogen fixation across the tropical oligotrophic regions in the model. Through the comparison between calibrated models, this thesis quantifies the advantages of including iron in a global biogeochemistry model and reveals how important iron is for future nitrogen fixation trends. It furthermore illustrates the interconnection between tropical upwelling and oligotrophic regions.

Description: Results of the of the present study provide a strong indication that reproductive periods of the bladderwrack Fucus vesiculosus is tuned by environmental conditions, such as day length, although it cannot be entirely ruled out that genetic constitution may play a role, as well. Furthermore results of the present study identified high temperatures as the most challenging condition for alga recruitment. Sea surface temperature rise could therefore be one of the reasons for the decline of F. vesiculosus populations in the Baltic Sea over the last few decades, particularly in the marginal environments (〈 7 psu). Additionally, fertility of F. vesiculosus from the marginal region, in contrast to all other regions, was very low, which also indicates towards a lower capacity to deal with environmental changes. A rather high germination success of some sibling groups (F. vesiculosus) under various environmental conditions, however, is promising in the light of adaptation to climate change.

Description: Oceanic emissions of sulfur containing trace gases alter global atmospheric chemistry. The gases act for example as aerosol precursors and change the radiative budget of the Earth, with a significant impact on climate. Large uncertainties exist in the amount of sulfur gases emitted from the ocean, and a gap in the atmospheric budget of carbonyl sulfide – the most abundant sulfur gas in the atmosphere – has been suggested to result from tropical ocean emissions. This thesis uses new shipbased measurements from the tropical Pacific and Indian Ocean together with models to quantify these emissions. Three studies were performed: 1) A 3D model study to test how oceanic emissions can be represented in atmospheric chemistry climate models, 2) A combination of new shipbased data and box model calculation to derive a global emission estimate of carbonyl sulfide and 3) a detailed process study of production processes and their drivers for the gases carbonyl sulfide and carbon disulfide in the Eastern tropical South Pacific. Together, the results yield a new temporally and spatially resolved emission climatology of the three gases.

Description: Small to meso-scale distribution of Baltic cod (Gadus morhua L.) as resolved by hydroacoustics: Habitat preferences, environmental limits, and resulting implications for stock development

Description: Over the past decade large carbonate mound structures, situated in several provinces along the European continental margin have been discovered in 600 to 1200 m water depths. These structures appear as single, conical or ridge-like features, which are often colonised by a deep-water coral ecosystem. Cold-water corals Lophelia pertusa and Madrepora oculata dominate the coral community. Associated sessile and vagile fauna, like sponges, soft corals, bivalves, gastrodpods, crinoids, brachiopods, and fish make these bioherms as diverse as their tropical counterparts. The objective of this study is to reconstruct past environmental settings locked in the sediments of a carbonate mound in the northern Porcupine Seabight, west off Ireland. Detailed investigations were concentrated on seven 3.5 to 6 m long sediment cores, retrieved from Propeller Mound and its closer vicinity. This mound is a ~150 m high structure within the Hovland Mound province, a cluster of several sea-floor protruding carbonate mounds. A first detailed analysis on benthic foraminiferal assemblages in the vicinity of a carbonate mound was performed on two sediment cores, one located on top of the mound and a second one from an off-mound position further north as control site. The off-mound samples reveal two different assemblages: (1) an Interglacial group dominated by infaunal species reflecting present-day environmental and oceanographic conditions with high nutrient flux to the sea-floor and low sediment accumulation under a strong hydrodynamic regime, and (2) a Glacial group, dominated by cassidulinid species, which describe an influence of polar conditions with low nutrient supply and cold intermediate waters. A single species, Elphidium excavatum, dominates the lower core section and is described here as a species displaced from shallow shelf areas. This removal of sediments from the shelf is related to a first advance of the Irish Ice Sheet onto the Irish Mainland shelf, inducing high sedimentation rates of ~28 cm/kyr with a coeval sea-level drop of ~50 m during Late Marine Isotope Stage (MIS) 3. Due to an incomplete stratigraphic record in the on-mound core, the same species of the off-mound assemblages have been grouped in on-mound samples. The results indicate a dominance of the Interglacial group, whereas the Glacial group is less abundant throughout the entire core. This pattern indicates the lack of glacial time intervals in the on-mound core, which is coherent with stable oxygen isotope data and U/Th dates on coral fragments, presenting only interglacial/-stadial values and ages. A third assemblage is abundant in samples of the on-mound core showing elevated epibenthic species not occurring in offmound samples or only to a minor degree. This Mound group shows a great affinity to strong currents, high nutrient availability and is supposed to indicate Mediterranean Outflow Water in the northern Porcupine Seabight, as well as a higher coral cover on Propeller Mound in an earlier interglacial period. A Late Pleistocene decline in mound growth for Propeller Mound is suggested by a decrease of the Mound group towards the Holocene, which might face its complete burial in the future as this already happened to the buried mounds of the Magellan Mound province further north. Detailed information on off-mound sediment structures and contents from visual core description and the interpretation of Computer Tomographic images were used to evaluate sedimentary processes in glacial and interglacial periods. The sediments portray the depositional history of the past ~31 kyr BP, mainly controlled by sea-level fluctuations and the climate regime with the advance and retreat of the Irish Ice Sheet onto the Irish Mainland Shelf. A first advance of glaciers is indicated by a turbiditic release slightly older than 31 kyr BP, coherent with Heinrich event 3. During MIS 3 and MIS 2 shelf erosion prevailed with abundant gravity flows and turbidity currents. A change from glaciomarine to hemipelagic contourite sedimentation during the onset of the Holocene indicates the establishment of the strong, present-day hydrodynamic regime at intermediate depths. A general decrease in sediment accumulation with decreasing distance towards Propeller Mound was discovered throughout the entire core sections. This suggests that currents (turbidity currents, gravity flows, bottom currents) have had a strong impact on sediment accumulation at the mound base for the past ~31 kyr BP. Finally, the reconstructed environmental setting deduced from sedimentary and micropaleontological analyses portrays the boundary conditions of the habitable range for the cold-water coral Lophelia pertusa. The growth of this ecosystem occurs during interglacial and interstadial periods, whereas a glacial retreat of corals is documented in the absence of glacial sediments in the onmound core. These conclusions are summarised in a model – the Mound Factory – which efficiently accounts for the moundd development covering the period of the Northern Hemisphere Glaciation (past 3.1–2.5 Ma).

Description: Fouling communities are distributed worldwide, particularly in shallow and exposed marine systems. Due to their fast growth and their capacity to settle on artificial surfaces, these assemblages represent a suitable study system for ecologists and are therefore often used to investigate ecological models. Investigating broad scale ecological models has been of key interest since the time of the famous naturalist Charles Darwin. Two examples of well recognized macro-ecological patterns are i) the ‘latitudinal gradient of species richness’ and ii) the ‘relationship between local and regional diversity’. The first is the oldest paradigm about a large-scale ecological pattern and simply states that the tropical regions are richer in species than temperate and polar regions at higher latitudes. However, when exploring global diversity patterns, it is essential to comprehend not only the importance of differences in spatial scale for the observed patterns, but also how diversity at one scale may relate to diversity at a different scale. Indeed, several studies have shown that the number of species within small localities may increase either linearly or asymptotically with regional species richness (relationship between local and regional diversity). Exploring global diversity patterns still constitutes an important challenge for ecologists because it reflects the need to determine the current status of biodiversity. With the knowledge of today’s biodiversity, we can predict its future status and, as a consequence, be able to provide new answers to its probable response to phenomena such as climate change. In the present work, I investigate global diversity patterns in marine fouling communities, mainly by examining the previously mentioned macro-ecological patterns, i.e. the latitudinal gradient of species richness and the relationship between local and regional diversity. Previous studies investigating the local-regional diversity relationship have often assessed the number of species in a region by consulting available species lists. However, regional species pools based on such inventories may include species not susceptible to recruit into the community considered because they are restricted to different habitats and seasons. With the purpose of dealing with these difficulties, a few investigations have estimated regional richness based on local samples but confirmed a strong bias in relation to sampling effort. In order to optimize the quality of regional richness estimations, the current study develops a new statistical tool for estimating regional richness based on a limited number of replicates. Using three data-sets with a large number of replicates from different temperate shallow water habitats, I compare common richness estimators against the asymptote of the species accumulation curve, which was used as a reference for true regional richness. Subsequently and more importantly, the mis-estimation was quantified as a function of sampling effort. To complete this work, the relationship between local and regional diversity was expanded by integrating two categories of diversity (taxonomic and functional) and different successional stages at two different scales: European and global. At the European scale, the shape of the pattern was compared for different methods in assessing regional richness: species colonizing during a given period (transient regional richness) versus species colonizing during any phase of the experiment (total regional richness). At the global scale, I further examine whether the diversity of local communities is affected by parameters other than regional richness, such as number of functional groups or availability of resources. The results of this investigation support a clear influence of latitude on local species richness in marine fouling communities. In Chapter I, I show that tropical regions hold more marine fouling species when compared to areas at higher latitudes. In what concerns the regional richness assessment, I conclude that regional richness can be estimated based on a limited number of samples and that the quality of the estimation increases with sample effort. Moreover, the strength of the inevitable mis-estimation can be quantified (Chapter II). In addition, at the European scale (Chapter III), it was found that the shape of the relationship between local and regional diversity is sensitive to successional stage, the way regional richness is estimated and the dimension of diversity considered. The relevant regional richness, i.e. the regionally available colonizers, seems to vary in time and is larger when pooling all sampling events. As a consequence, the relationship between local and regional diversity is also influenced by the method in which regional richness is estimated. At a global scale (Chapter IV), the relationship between local and regional diversity in fouling assemblages is also affected by the succession process, if either taxonomic or functional diversity are considered. Local taxonomic diversity exhibits saturation at early stages of succession while saturation of local functional richness occurs later. In addition, functional groups were reported as the most influential predictor for local species richness.