Spatial and temporal variation in fish populations and assemblages in coastal waters of the northern Baltic Proper

The distribution and traits of fish are of interest both ecologically and socio-economically. In this thesis, phenotypic and structural variation in fish populations and assemblages was studied on multiple spatial and temporal scales in shallow coastal areas in the archipelago of the northern Baltic Proper. In Lumparn basin in Åland Islands, the fish assemblage displayed significant seasonal variation in depth zone distribution. The results indicate that investigating both spatial and temporal variation in small scale is crucial for understanding patterns in fish distribution and community structure in large scale. The local population of Eurasian perch Perca fluviatilis L displayed habitat-specific morphological and dietary variation. Perch in the pelagic zone were on average deeper in their body shape than the littoral ones and fed on fish and benthic invertebrates. The results differ from previous studies conducted in freshwater habitats, where the pelagic perch typically are streamlined in body shape and zooplanktivorous. Stable isotopes of carbon and nitrogen differed between perch with different stomach contents, suggesting differentiation of individual diet preferences. In the study areas Lumparn and Ivarskärsfjärden in Åland Islands and Galtfjärden in Swedish east coast, the development in fish assemblages during the 2000’s indicated a general shift towards higher abundances of small-bodied lower-order consumers, especially cyprinids. For European pikeperch Sander lucioperca L., recent declines in adult fish abundances and high mortalities (Z = 1.06–1.16) were observed, which suggests unsustainably high fishing pressure on pikeperch. Based on the results it can be hypothesized that fishing has reduced the abundances of large predatory fish, which together with bottom-up forcing by eutrophication has allowed the lower-order consumer species to increase in abundances. This thesis contributes to the scientific understanding of aquatic ecosystems with new descriptions on morphological and dietary adaptations in perch in brackish water, and on the seasonal variation in small-scale spatial fish distribution. The results also demonstrate anthropogenic effects on coastal fish communities and underline the urgency of further reducing nutrient inputs and regulating fisheries in the Baltic Sea region.

Distribution and role of macrophytes in coastal lagoons : implications of critical shifts

Crossroads, crucibles and refuges are three words that may describe natural coastal lagoon environments. The words refer to the complex mix of marine and terrestrial influences, prolonged dilution due to the semi-enclosed nature and the function of a habitat for highly diverse plant and animal communities, some of which are endangered. To attain a realistic picture of the present situation, high vulnerability to anthropogenic impact should be added to the description. As the sea floor in coastal lagoons is usually entirely photic, macrophyte primary production is accentuated compared with open sea environments. There is, however, a lack of proper knowledge on the importance of vegetation for the general functioning of coastal lagoon ecosystems. The aim of this thesis is to assess the role of macrophyte diversity, cover and species identity over temporal and spatial scales for lagoon functions, and to determine which steering factors primarily restrict the qualitative and quantitative composition of vegetation in coastal lagoons. The results are linked to patterns of related trophic levels and the indicative potential of vegetation for assessment of general conditions in coastal lagoons is evaluated. This thesis includes five field studies conducted in flads and glo-flads in the brackish water northern Baltic Sea. Flads and glo-flads are defined as a Baltic variety of coastal lagoons, which due to an inlet threshold and post-glacial landuplift slowly will be isolated from the open sea. This process shrinks inlet size, increases exposure and water retention, and is called habitat isolation. The studied coastal lagoons are situated in the archipelago areas of the eastern coast of Sweden, the Åland Islands and the south-west mainland of Finland, where land-uplift amounts to ca. 5 mm/ per year. Out of 400 evaluated sites, a total of 70 lagoons varying in inlet size, archipelago position and anthropogenic influence to cover for essential environmental variation were chosen for further inventory. Vegetation composition, cover and richness were measured together with several hydrographic and morphometric variables in the lagoons both seasonally and inter-annually to cover for general regional, local and temporal patterns influencing lagoon and vegetation development. On smaller species-level scale, the effects of macrophyte species identity and richness for the fish habitat function were studied by examining the influence of plant interaction on juvenile fish diversity. Thus, the active election of plant monoand polycultures by fish and the diversity of fish in the respective culture were examined and related to plant height and water depth. The lagoons and vegetation composition were found to experience a regime shift initiated by increased habitat isolation along with land-uplift. Vegetation composition altered, richness decreased and cover increased forming a less isolated and more isolated regime, named the vascular plant regime and charophyte regime, respectively according to the dominant vegetation. As total phosphorus in the water, turbidity and the impact of regional influences decreased in parallel, the dominance of charophytes and increasing cover seemed to buffer and stabilize conditions in the charophyte regime and indicated an increased functional role of vegetation for the lagoon ecosystem. The regime pattern was unaffected by geographical differences, while strong anthropogenic impact seemed to distort the pattern due to loss of especially Chara tomentosa L. in the charophyte regime. The regimes were further found unperturbed by short-time temporal fluctuations. In fact the seasonal and inter-annual dynamics reinforced the functional difference between the regimes by the increasing role of vegetation along habitat isolation and the resemblance to lake environments for the charophyte regime. For instance, greater total phosphorus and chlorophyll a concentrations in the water in the beginning of the season in the charophyte regime compared with the vascular plant regime presented a steeper reduction to even lower values than in the vascular plant regime along the season. Despite a regional importance and positive relationship of macrophyte diversity in relation to trophic diversity, species identity was underlined in the results of this thesis, especially with decreasing spatial scale. This result was supported partly by the increased role of charophytes in the functioning of the charophyte regime, but even more explicitly by the species-specific preference of juvenile fish for tall macrophyte monocultures. On a smaller species-level scale, tall plant species in monoculture seemed to be able to increase their length, indicating that negative selection forms preferred habitat structures, which increase fish diversity. This negative relationship between plant and fish diversity suggest a shift in diversity patterns among trohic levels on smaller scale. Thus, as diversity patterns seem complex and diverge among spatial scales, it might be ambiguous to extend the understanding of diversity relationships from one trophic level to the other. All together, the regime shift described here presents similarities to the regime development in marine lagoon environments and shallow lakes subjected to nutrient enrichment. However, due to nutrient buffering by vegetation with increased isolation and water retention as a consequence of the inlet threshold, the development seems opposite to the course along an eutrophication gradient described in marine lagoons lacking an inlet threshold, where the role of vegetation decreases. Thus, the results imply devastating consequences of inlet dredging (decreasing isolation) in terms of vegetation loss and nutrient release, and call for increased conservational supervision. Especially the red listed charophytes would suffer negatively from such interference and the consequences are likely to also deteriorate juvenile fish production. The fact that a new species to Finland, Chara connivens Salzm. Ex. Braun 1835 was discovered during this study further indicates a potential of the lagoons serving as refuges for rare species.

Streptococcus agalactiae (GBS) PNA FISH -tekniikan soveltuminen B-ryhmän streptokokin tunnistukseen

B-ryhmän beetahemolyyttinen streptokokki (GBS = Group B Streptococcus, Streptococcus agalactiae)aiheuttaa vakavia infektioita yleensä astasyntyneillä. Tartunta saadaan yleensä synnytyskanavasta ja riskitekijöinä ovat muun muassa keskosuus, ennenaikainen lapsivedenmeno ja äidin runsas Bstreptokokkikolonisaatio emättimessä. Bakteerin tunnistukseen käytetään tällä hetkellä viljelytekniikkaa, jonka tulos saadaan vasta 24-48 tunnin kuluttua. Opinnäytetyöni tarkoituksena on tutkia uutta ja nopeampaa tunnistusmenetelmää: GBS PNA FISH - tekniikkaa (Peptide Nucleic Acid Fluorescence in Situ Hybridization). Tarkoituksena on tutkia tekniikan spesifiteettiä ja sensitiviteettiä. Tekniikan spesifiteettiä tutkitaan B-ryhmän beetahemolyyttisellä streptokokilla sekä kuudella muulla emättimen normaaliflooraan kuuluvalla bakteerilajilla. Yhteensä bakteerikantoja on tutkimuksessa mukana 48 kappaletta. Tämän lisäksi tutkitaan myös tekniikan sensitiviteettiä, jota tutkitaan bakteereista tehdyn laimennossarjan avulla. Sensitiviteetti tutkitaan bakteeriseoksesta, jonne on B-ryhmän beetahemolyyttisen streptokokin lisäksi lisätty muita emättimen normaaliflooran bakteereita. Lisäksi sensitiviteetti tutkitaan pelkällä B-ryhmän beetahemolyyttisellä streptokokilla käyttäen sekä normaalia että bakteerin rikastusmenetelmää. Testeistä saadut tulokset tulkitaan fluoresenssimikroskoopin avulla. GBS PNA FISH -tekniikan spesifiteetti todettiin erittäin hyväksi. Tekniikka tunnisti kaikki B-ryhmän beetahemolyyttiset streptokokit positiivisiksi ja kaikki muut lajit antoivat negatiivisen tuloksen. B-streptokokin positiivisuus oli erotettavissa mikroskopoitaessa vahvana fluoresointina, kun taas muut lajit eivät fluoresoineet lainkaan. GBS PNA FISH -tekniikan sensitiivisyyden tulokset eivät kuitenkaan täyttäneet odotuksia. Ainoastaan bakteerin rikastusmenetelmällä saadut tulokset olivat loistavia, mutta bakteeriseoksella ja pelkällä B-ryhmän beetahemolyyttisellä streptokokilla saadut tulokset olivat lähes olemattomia. Rikastusmenetelmän kaikki laimennokset fluoresoivat positiivisina, kun taas muissa tapauksissa vain vahvin liuos antoi jonkinlaista positiivista fluoresointia. GBS PNA FISH -tekniikan spesifiteetti todettiin hyväksi. Tekniikan sensitiviteetti ei kuitenkaan vastaa käyttötarkoitusta ja todellisessa tilanteessa tekniikka ei pystyisi tunnistamaan sille spesifistä bakteeria muiden bakteerien joukosta.

Spatiotemporal Aspects of Environmental Monitoring in the Complex Coastal Region of Southwest Finland

The management and conservation of coastal waters in the Baltic is challenged by a number of complex environmental problems, including eutrophication and habitat degradation. Demands for a more holistic, integrated and adaptive framework of ecosystem-based management emphasize the importance of appropriate information on the status and changes of the aquatic ecosystems. The thesis focuses on the spatiotemporal aspects of environmental monitoring in the extensive and geomorphologically complex coastal region of SW Finland, where the acquisition of spatially and temporally representative monitoring data is inherently challenging. Furthermore, the region is subject to multiple human interests and uses. A holistic geographical approach is emphasized, as it is ultimately the physical conditions that set the frame for any human activity. Characteristics of the coastal environment were examined using water quality data from the database of the Finnish environmental administration and Landsat TM/ETM+ images. A basic feature of the complex aquatic environment in the Archipelago Sea is its high spatial and temporal variability; this foregrounds the importance of geographical information as a basis of environmental assessments. While evidence of a consistent water turbidity pattern was observed, the coastal hydrodynamic realm is also characterized by high spatial and temporal variability. It is therefore also crucial to consider the spatial and temporal representativeness of field monitoring data. Remote sensing may facilitate evaluation of hydrodynamic conditions in the coastal region and the spatial extrapolation of in situ data despite their restrictions. Additionally, remotely sensed images can be used in the mapping of many of those coastal habitats that need to be considered in environmental management. With regard to surface water monitoring, only a small fraction of the currently available data stored in the Hertta-PIVET register can be used effectively in scientific studies and environmental assessments. Long-term consistent data collection from established sampling stations should be emphasized but research-type seasonal assessments producing abundant data should also be encouraged. Thus a more comprehensive coordination of field work efforts is called for. The integration of remote sensing and various field measurement techniques would be especially useful in the complex coastal waters. The integration and development of monitoring system in Finnish coastal areas also requires further scientific assesement of monitoring practices. A holistic approach to the gathering and management of environmental monitoring data could be a cost-effective way of serving a multitude of information needs, and would fit the holistic, ecosystem-based management regimes that are currently being strongly promoted in Europe.

Assessment of Coastal Bird Populations and Habitats on the Finnish Coast of the Baltic Sea: Implications for Monitoring and Management

Coastal birds are an integral part of coastal ecosystems, which nowadays are subject to severe environmental pressures. Effective measures for the management and conservation of seabirds and their habitats call for insight into their population processes and the factors affecting their distribution and abundance. Central to national and international management and conservation measures is the availability of accurate data and information on bird populations, as well as on environmental trends and on measures taken to solve environmental problems. In this thesis I address different aspects of the occurrence, abundance, population trends and breeding success of waterbirds breeding on the Finnish coast of the Baltic Sea, and discuss the implications of the results for seabird monitoring, management and conservation. In addition, I assess the position and prospects of coastal bird monitoring data, in the processing and dissemination of biodiversity data and information in accordance with the Convention on Biological Diversity (CBD) and other national and international commitments. I show that important factors for seabird habitat selection are island area and elevation, water depth, shore openness, and the composition of island cover habitats. Habitat preferences are species-specific, with certain similarities within species groups. The occurrence of the colonial Arctic Tern (Sterna paradisaea) is partly affected by different habitat characteristics than its abundance. Using long-term bird monitoring data, I show that eutrophication and winter severity have reduced the populations of several Finnish seabird species. A major demographic factor through which environmental changes influence bird populations is breeding success. Breeding success can function as a more rapid indicator of sublethal environmental impacts than population trends, particularly for long-lived and slowbreeding species, and should therefore be included in coastal bird monitoring schemes. Among my target species, local breeding success can be shown to affect the populations of the Mallard (Anas platyrhynchos), the Eider (Somateria mollissima) and the Goosander (Mergus merganser) after a time lag corresponding to their species-specific recruitment age. For some of the target species, the number of individuals in late summer can be used as an easier and more cost-effective indicator of breeding success than brood counts. My results highlight that the interpretation and application of habitat and population studies require solid background knowledge of the ecology of the target species. In addition, the special characteristics of coastal birds, their habitats, and coastal bird monitoring data have to be considered in the assessment of their distribution and population trends. According to the results, the relationships between the occurrence, abundance and population trends of coastal birds and environmental factors can be quantitatively assessed using multivariate modelling and model selection. Spatial data sets widely available in Finland can be utilised in the calculation of several variables that are relevant to the habitat selection of Finnish coastal species. Concerning some habitat characteristics field work is still required, due to a lack of remotely sensed data or the low resolution of readily available data in relation to the fine scale of the habitat patches in the archipelago. While long-term data sets exist for water quality and weather, the lack of data concerning for instance the food resources of birds hampers more detailed studies of environmental effects on bird populations. Intensive studies of coastal bird species in different archipelago areas should be encouraged. The provision and free delivery of high-quality coastal data concerning bird populations and their habitats would greatly increase the capability of ecological modelling, as well as the management and conservation of coastal environments and communities. International initiatives that promote open spatial data infrastructures and sharing are therefore highly regarded. To function effectively, international information networks, such as the biodiversity Clearing House Mechanism (CHM) under the CBD, need to be rooted at regional and local levels. Attention should also be paid to the processing of data for higher levels of the information hierarchy, so that data are synthesized and developed into high-quality knowledge applicable to management and conservation.

Natural Science in the Bothnian Sea During 1975-2008: A Review

The Baltic Sea is one of the most studied areas in the world. However, parts of its northernmost reach, the Bothnian Sea, seem to be under represented in the natural scientific literature compared to other parts of the Baltic. The Bothnian Sea represents a unique inland sea environment for the scientific community to study due to its shallowness and low salinity. The natural sciences research carried out on the Bothnian Sea has been reviewed between 1975 and 2008. This time period was chosen to continue on from an earlier review paper ending in 1974. Along with the number of papers published the goal was also to review the content of the papers, indentifying dominating themes to evaluate gaps in the current knowledge on the Bothnian Sea and provide recommendations for topics of future research focus. In a classification into specific research topics biodiversity was the leading research focus followed by chemical and physical oceanography, pollution and toxins, and fish biology and fisheries. The current good condition of the Bothnian Sea is highly valued with its historically less eutrophic and clearer waters when compared to the Baltic. However, today the Bothnian Sea is facing eutrophication resulting from nutrient-rich water transported by the many rivers draining into it from Sweden and Finland making it an area in need of protection and preservation. More human activity will also concentrate on the Bothnian Sea in the future. Therefore the use of the sea and its coastal areas must be planned carefully to minimize the harmful effects of this increasing human activity. To achieve this more information is needed for the basis of Integrated Coastal Zone Management (ICZM) and maritime spatial planning (MSP). For example, for the Bothnian Sea the information on the underwater nature which is essential for ICZM is so far missing to a large extent. Specific biological, chemical and physical oceanographic information is needed to combine with economic analyses and environmental policies regarding this region. More research of a multidisciplinary nature is required on the unique Bothnian Sea environment and this we feel is best achieved through a joint Finnish-Swedish research strategy.

Fish ja periaatteiden tyhjyys

Kirjallisuusarvostelu

Identification and application of bile metabolites to assess the exposure of fish to pharmaceuticals in the environment

Thousands of tons of pharmaceuticals are consumed yearly worldwide. Due to the continuous and increasing consumption and their incomplete elimination in wastewater treatment plants (WWTP), pharmaceuticals and their metabolites can be detected in receiving waters, although at low concentrations (ng to low μg/L). As bioactive molecules the presence of pharmaceuticals in the aquatic environment must be considered potentially hazardous for the aquatic organisms. In this thesis, the biotransformation and excretion of pharmaceuticals in fish was studied. The main biotransformation pathways of three anti‐inflammatory drugs, diclofenac, naproxen and ibuprofen, in rainbow trout were glucuronidation and taurine conjugation of the parent compounds and their phase I metabolites. The same metabolites were present in fish bile in aquatic exposures as in fish dosed with intraperitoneal injection. Higher bioconcentration factor in bile (BCFbile) was found for ibuprofen when compared to diclofenac and naproxen. Laboratory exposure studies were followed by a study of uptake of pharmaceuticals in a wild fish population living in lake contaminated with WWTP effluents. Of the analyzed 17 pharmaceuticals and six phase I metabolites, only diclofenac, naproxen and ibuprofen was present in bream and roach bile. It was shown, that diclofenac, naproxen and ibuprofen excreted by the liver can be found in rainbow trout and in two native fish species living in the receiving waters. In the bream and roach bile, the concentrations of diclofenac, naproxen and ibuprofen were roughly 1000 times higher than those found in the lake water, while in the laboratory exposures, the bioconcentration of the compounds and their metabolites in rainbow trout bile were at the same level as in wild fish or an order of magnitude higher. Thus, the parent compounds and their metabolites in fish bile can be used as a reliable biomarker to monitor the exposure of fish to environmental pharmaceuticals present in water receiving discharges from WWTPs.

Interpreting marine benthic ecosystem functioning in coastal waters: validating the biological trait concept

Coastal areas harbour high biodiversity, but are simultaneously affected by rapid degradations of species and habitats due to human interactions. Such alterations also affect the functioning of the ecosystem, which is primarily governed by the characteristics or traits expressed by the organisms present. Marine benthic fauna is nvolved in numerous functions such as organic matter transformation and transport, secondary production, oxygen transport as well as nutrient cycling. Approaches utilising the variety of faunal traits to assess benthic community functioning have rapidly increased and shown the need for further development of the concept. In this thesis, I applied biological trait analysis that allows for assessments of a multitude of categorical traits and thus evaluation of multiple functional aspects simultaneously. I determined the functional trait structure, diversity and variability of coastal zoobenthic communities in the Baltic Sea. The measures were related to recruitment processes, habitat heterogeneity, large-scale environmental and taxonomic gradients as well as anthropogenic impacts. The studies comprised spatial scales from metres to thousands of kilometres, and temporal scales spanning one season as well as a decade. The benthic functional structure was found to vary within and between seagrass landscape microhabitats and four different habitats within a coastal bay, in papers I and II respectively. Expressions of trait categories varied within habitats, while the density of individuals was found to drive the functional differences between habitats. The findings in paper III unveiled high trait richness of Finnish coastal benthos (25 traits and 102 cateogries) although this differed between areas high and low in salinity and human pressure. In paper IV, the natural reduction in taxonomic richness across the Baltic Sea led to an overall reduction in function. However, functional richness in terms of number of trait categories remained comparatively high at low taxon richness. Changes in number of taxa within trait categories were also subtle and some individual categories were maintained or even increased. The temporal analysis in papers I and III highlighted generalities in trait expressions and dominant trait categories in a seagrass landscape as well as a “type organism” for the northern Baltic Sea. Some initial findings were made in all four papers on the role of common and rare species and traits for benthic community functioning. The findings show that common and rare species may not always express the same trait categories in relation to each other. Rare species in general did not express unique functional properties. In order to advance the understanding of the approach, I also assessed some issues concerning the limitations of the concept. This was conducted by evaluating the link between trait category and taxonomic richness using especially univariate measures. My results also show the need to collaborate nationally and internationally on safeguarding the utility of taxonomic and trait data. The findings also highlight the importance of including functional trait information into current efforts in marine spatial planning and biomonitoring.