Environmental factors influence species and habitats on multiple scales creating a mosaic of distribution patterns. Studying factors shaping these patterns are central to our understanding of population dynamics and ultimately ecosystem functioning. Information on the distribution of resources and conservation values are also highly needed in marine management as coastal areas are increasingly influenced by human activities. In this thesis, large-scale field data is used to explore how strong environmental gradients found on multiple scales in the coastal areas of the Baltic Sea influence fish habitats. The underlying concepts are based in the field of species distribution modelling, whereby habitat maps can be produced using environmental layers in a geographic information system. Distribution modelling is further used to address both ecological and applied questions by examining effects of habitat limitation on fish population sizes and to evaluate management actions aimed at habitat conservation. I show that specific habitat requirements for fish species of both freshwater and marine origin can be described using environmental variables and that species-environment relationships can be used to predict the distribution of early life-stages of fish in the Baltic Sea archipelagos. Further, predicted habitat availability of a specific life-stage was directly related to adult population size of Eurasian perch Perca fluviatilis, signifying that the abundance of large predatory fish can be limited by specific recruitment habitats. Lastly, by predicting the distribution of an assemblage of coastal fish species and their associated habitats, an assessment of a network of marine protected areas was performed. Results revealed large gaps in the current network and identified areas suitable for future protection. By demonstrating how current habitat protection can be improved by including critical habitats for coastal fish population sizes this thesis points to the benefits of integrating nature conservation and fisheries management. Based on these findings I conclude that species distribution modelling provides a suitable analytical framework for assessing the habitat requirements of organisms. An increased understanding of habitat-population relationships and an ability to accurately map ecologically important features will be of great value for an ecosystem-based marine management. ­ / Felaktigt tryckt som Digital Comprehensive Summaries of Uppsala Dissertations from the Faculty of Science and Technology 709

In boreal catchments, stream water chemistry is influenced and controlled by several landscape factors. The influence of spatially distributed variables is in turn dependent on the hydrological scale. Headwater streams have larger variability of water chemistry, and thus together represent a large biodiversity, and therefore need to be monitored in official environmental assessments. One objective of this study was, using Geographical Information Systems (GIS), to analyse co-variation between landscape variables and water chemistry and to determine which of the landscape variables have a major influence on the concentration of dissolved organic carbon (DOC) in headwater streams. Another objective was to find a simple method for predicting sources of DOC, using official map data and publically available GIS applications. Totally 85 headwater catchments (0.1-4 km2) in the county of Värmland, western south Sweden, were used in the study. Water chemistry was analysed for water sampled at low, medium and high flows, and landscape variables were extracted from official map data sources: topographic maps, a digital elevation model (DEM, 50 m grid), and vegetation data. Statistical analyses showed that topography (mean slope and mean topographic wetness index (TWI)) and wetland cover often correlated well with DOC in headwater catchments. Official map data could satisfactorily extract landscape variables (mean slope, mean TWI) that were useful in predicting stream water chemistry (DOC). A high-resolution elevation model, which was generated by interpolation of photogrammetric data, was used to calculate and evaluate two different wetness indices and their ability to predict the occurrence of wetlands in six catchments of different sizes and topography. The SAGA (System for Automated Geoscientific Analyses) wetness index (SWI) gave substantially better results than the TWI. The effects of resolution of DEMs on calculations of the SWI were investigated using 5, 10, 25 and 50 m grids. The results showed that SWI values increased with increasing cell size. The near linear increment of mean values for resolutions 10-50 m suggests a independence of terrain type and catchment size, which supported previous findings that indicated that mean slope and mean wetness index calculated from coarse elevation models may be used for prediction of DOC in headwater streams.

The trophic state of lakes is commonly defined by the concentration of nutrients in the water column. High nutrient concentrations generate high phytoplankton production, and lakes with low nutrient concentrations are considered low-productive. This simplified view of lake productivity ignores the fact that benthic primary producers and heterotrophic bacteria can be important basal producers in lake ecosystems. In this thesis I have studied clear-water and brown-water lakes with respect to primary production, respiration and bacterial production based on allochthonous organic carbon. These processes were quantified in pelagic and benthic habitats on temporal and spatial scales. I also calculated the net ecosystem production of the lakes, defined as the difference between gross primary production (GPP) and respiration (R). The net ecosystem production indicates whether a lake is net heterotrophic (GPP < R), net autotrophic (GPP > R) or in metabolic balance (GPP = R). Net heterotrophic lakes are sources of carbon dioxide (CO2) to the atmosphere since respiration in these lakes, by definition, is subsidized by an external organic carbon source. External organic carbon is transported to lakes from the terrestrial environment via inlets, and can serve as a carbon source for bacteria but it also limits light availability for primary producers by absorbing light. On a seasonal scale, four of the clear-water lakes studied in this thesis were dominated by primary production in the soft-bottom benthic habitat and by respiration in the pelagic habitat. Concentrations of dissolved organic carbon (DOC) were low in the lakes, but still high enough to cause the lakes to be net heterotrophic. However, the lakes were not low-productive due to the high production in the benthic habitat. One of the clear-water lakes was studied also during the winter and much of the respiration under ice was supported by the benthic primary production from the previous summer. This is in contrast to brown-water lakes where winter respiration is suggested to be supported by allochthonous organic carbon. By studying lakes in a DOC gradient (i.e. from clear-water to brown-water lakes) I could draw two major conclusions. The lakes became less productive since benthic primary production decreased with increasing light extinction, and the lakes became larger sources of CO2 to the atmosphere since pelagic respiration was subsidized by allochthonous organic carbon. Thus, lake carbon metabolism can have an important role in the global carbon cycle due to their processing of terrestrial organic carbon and to their possible feedback effects on the climate system.

Cross-boundary movements of energy and material are ubiquitous. Freshwater ecosystems receive nutrients, dissolved, and particulate organic matter from adjacent terrestrial ecosystems, whereas terrestrial ecosystems mainly receive prey organisms and detritus deposited by physical processes such as floods from freshwater ecosystems. Within lakes, fish are considered as integrators between habitats due to their high mobility, although they often occupy either near-shore littoral or open-water pelagic habitats and develop habitat-specific morphologies. Such intra-population divergence in morphological traits might limit the use of multiple habitats. In this thesis, I first focused on quantity and quality of reciprocal fluxes of particulate organic matter between freshwater and terrestrial ecosystems and responses of recipient consumers. Freshwater ecosystems generally received higher amounts of externally-produced resources than terrestrial ecosystems. Despite this discrepancy, aquatic and terrestrial consumer responses were similar, likely due to the differences in resource quality. Second, I investigated the potential of particulate organic carbon (POC) supporting benthic food webs in lakes; a pathway that has largely been neglected in previous studies. I found that POC can substantially subsidize the benthic food web and that the effects on the benthic food web were transferred to the pelagic habitat, thus emphasizing the importance of benthic pathways for pelagic production. Third, I examined how water transparency can affect intra-population divergence in perch (Perca fluviatilis). I observed that increased water transparency can considerably increase morphological divergence between littoral and pelagic populations likely due to its effects on foraging. Finally, I investigated the effects of such intra-population divergence on littoral-pelagic food web coupling. I found that low morphological divergence corresponded with high overlap in resource use, whereas strong morphological divergence resulted in low overlap in resource use. Here littoral populations mainly utilized littoral resources and pelagic populations primarily utilized pelagic resources, indicating that habitat coupling might be strongly limited when intra-population divergence is high. In conclusion, although different ecosystems seem separated by distinct physical boundaries, these boundaries are often crossed. However, the development of habitat-specific adaptive traits might limit movement between apparently contiguous habitats.