Kirkkaasta sameaan : Meren kuormitus ja tila Saaristomerellä ja Ahvenanmaalla

Tässä raportissa tarkastellaan Saaristomeren ja Ahvenmaan merialueen kuormitusta ja tilaa ja niissä viime vuosikymmeninä tapahtuneita muutoksia vuosiin 2008 ja 2009 saakka. Pääpaino on ravinnekuormituksessa ja veden laadussa, mutta mukana on taustatietoa ja muun muassa katsaus kalakannoista. Saaristomeren ja Ahvenanmaan merialueen tila on heikentynyt huomattavasti viimeisten vuosikymmenten aikana. Erityisesti fosforin ja a-klorofyllin määrät ovat kasvaneet ja näkösyvyys pienenentynyt. Viimeisen kymmenen vuoden aikana pintaveden fosforipitoisuuden kasvu on laantunut suurimmassa osassa aluetta, mutta pohjanläheisessä vesikerroksessa ja talvella fosforipitoisuus on jatkanut kasvuaan monin paikoin viime vuosiin asti. Myös a-klorofyllin määrä on lisääntynyt ulko- ja välisaaristossa 2000-luvullakin. Viime vuosina klorofyllipitoisuuden kasvu on kuitenkin laantunut tai pitoisuus laskenut varsinkin rannikonläheisissä vesissä. Merialueen suurin paikallinen ravinnekuormittaja on maatalous, mutta varsinkin typpeä tulee runsaasti myös ilmalaskeumana. Sisäinen kuormitus ja virtausten mukana Suomenlahdelta ja muualta Itämereltä tulevat ravinteet heikentävät niin ikään merialueen tilaa. Muita ravinnelähteitä ovat mm. haja-asutus, jätevedenpuhdistamot ja kalankasvatus. Ahvenanmaan merialueelle ravinteita tulee enemmän muualta kuin Ahvenanmaalta. Hajakuormituksessa on tapahtunut viime vuosikymmeninä selviä vuodenaikaisia muutoksia. Vuosina 1990 – 2009 jokien mereen tuoma ravinne- ja kiintoainekuormitus on ollut talvisin ja syksyisin selvästi suurempaa kuin vuosina 1970 – 1989. Talviaikaisen kuormituksen lisääntyminen johtuu aiempaa lauhemmista talvista, ja antaa viitteitä ilmastonmuutoksen mukanaan tuomista muutoksista. Vaikka Saaristomeren ja Ahvenanmaan merialueen tilassa on ollut viime aikoina nähtävissä myönteisiäkin merkkejä, on niiden tila edelleen huolestuttava. Ainoa keino merialueen tilan parantamiseksi on kuormituksen huomattava vähentäminen. Vähentäminen koskee niin paikallisia ravinnelähteitä, erityisesti maataloutta, kuin muualta Itämereltä virtausten mukana tulevia ravinteita ja ilmalaskeumaa.

Of milquetoasts and daredevils : personalities in female eiders

Traditionally biologists have often considered individual differences in behaviour or physiology as a nuisance when investigating a population of individuals. These differences have mostly been dismissed as measurement errors or as non-adaptive variation around an adaptive population mean. Recent research, however, challenges this view. While long acknowledged in human personality studies, the importance of individual variation has recently entered into ecological and evolutionary studies in the form of animal personality. The concept of animal personality focuses on consistent differences within and between individuals in behavioural and physiological traits across time and contexts and its ecological and evolutionary consequences. Nevertheless, a satisfactory explanation for the existence of personality is still lacking. Although there is a growing number of explanatory theoretical models, there is still a lack of empirical studies on wild populations showing how traditional life-history tradeoffs can explain the maintenance of variation in personality traits. In this thesis, I first investigate the validity of variation in allostatic load or baseline corticosterone (CORT) concentrations as a measure for differences in individual quality. The association between CORT and quality has recently been summarised under the “CORT-fitness hypothesis”, which states that a general negative relationship between baseline CORT and fitness exists. I then continue to apply the concept of animal personality to depict how the life-history trade-off between survival and fecundity is mediated in incubating female eiders (Somateria mollissima), thereby maintaining variation in behaviour and physiology. To this end, I investigated breeding female eiders from a wild population that breeds in the archipelago around Tvärminne Zoological Station, SW Finland. The field data used was collected from 2008 to 2012. The overall aim of the thesis was to show how differences in personality and stress responsiveness are linked to a life-history context. In the four chapters I examine how the life-history trade-off between survival and fecundity could be resolved depending on consistent individual differences in escape behaviour, stress physiology, individual quality and nest-site selection. First, I corroborated the validity of the “CORT-fitness hypothesis”, by showing that reproductive success is generally negatively correlated with serum and faecal baseline CORT levels. The association between individual quality and baseline CORT is, however, context dependent. Poor body condition was associated with elevated serum baseline CORT only in older breeders, while a larger reproductive investment (clutch mass) was associated with elevated serum baseline CORT among females breeding late in the season. Interestingly, good body condition was associated with elevated faecal baseline CORT levels in late breeders. High faecal baseline CORT levels were positively related to high baseline body temperature, and breeders in poor condition showed an elevated baseline body temperature, but only on open islands. The relationship between stress physiology and individual quality is modulated by breeding experience and breeding phenology. Consequently, the context dependency highlights that this relationship has to be interpreted cautiously. Additionally, I verified if stress responsiveness is related to risk-taking behaviour. Females who took fewer risks (longer flight initiation distance) showed a stronger stress response (measured as an increase in CORT concentration after capture and handling of the bird). However, this association was modulated by breeding experience and body condition, with young breeders and those in poor body condition showing the strongest relationship between risktaking and stress responsiveness. Shy females (longer flight initiation distance) also incubated their clutch for a shorter time. Additionally, I demonstrated that stress responsiveness and predation risk interact with maternal investment and reproductive success. Under high risk of predation, females that incubated a larger clutch showed a stronger stress response. Surprisingly, these females also exhibited higher reproductive success than females with a weaker stress response. Again, these context dependent results suggest that the relationship between stress responsiveness and risk-taking behaviour should not be studied in isolation from individual quality and that stress responsiveness may show adaptive plasticity when individuals are exposed to different predation regimes. Finally, female risk-taking behaviour and stress coping styles were also related to nest-site choice. Less stress responsive females more frequently occupied nests with greater coverage that were farther away from the shoreline. Females nesting in nests with medium cover and farther from the shoreline had higher reproductive success. These results suggest that different personality types are distributed non-randomly in space. In this thesis I was able to demonstrate that personalities and stress coping strategies are persistent individual characteristics, which express measurable effects on fitness. This suggests that those traits are exposed to natural selection and thereby can evolve. Furthermore, individual variation in personality and stress coping strategy is linked to the alternative ways in which animals resolve essential life-history trade-offs.

Macroalgae across environmental gradients : tools for managing rocky coastal areas of the northern Baltic Sea

Macroalgae are the main primary producers of the temperate rocky shores providing a three-dimensional habitat, food and nursery grounds for many other species. During the past decades, the state of the coastal waters has deteriorated due to increasing human pressures, resulting in dramatic changes in coastal ecosystems, including macroalgal communities. To reverse the deterioration of the European seas, the EU has adopted the Water Framework Directive (WFD) and the Marine Strategy Framework Directive (MSFD), aiming at improved status of the coastal waters and the marine environment. Further, the Habitats Directive (HD) calls for the protection of important habitats and species (many of which are marine) and the Maritime Spatial Planning Directive for sustainability in the use of resources and human activities at sea and by the coasts. To efficiently protect important marine habitats and communities, we need knowledge on their spatial distribution. Ecological knowledge is also needed to assess the status of the marine areas by involving biological indicators, as required by the WFD and the MSFD; knowledge on how biota changes with human-induced pressures is essential, but to reliably assess change, we need also to know how biotic communities vary over natural environmental gradients. This is especially important in sea areas such as the Baltic Sea, where the natural environmental gradients create substantial differences in biota between areas. In this thesis, I studied the variation occurring in macroalgal communities across the environmental gradients of the northern Baltic Sea, including eutrophication induced changes. The aim was to produce knowledge to support the reliable use of macroalgae as indicators of ecological status of the marine areas and to test practical metrics that could potentially be used in status assessments. Further, the aim was to develop a methodology for mapping the HD Annex I habitat reefs, using the best available data on geology and bathymetry. The results showed that the large-scale variation in the macroalgal community composition of the northern Baltic Sea is largely driven by salinity and exposure. Exposure is important also on smaller spatial scales, affecting species occurrence, community structure and depth penetration of algae. Consequently, the natural variability complicates the use of macroalgae as indicators of human-induced changes. Of the studied indicators, the number of perennial algal species, the perennial cover, the fraction of annual algae, and the lower limit of occurrence of red and brown perennial algae showed potential as usable indicators of ecological status. However, the cumulated cover of algae, commonly used as an indicator in the fully marine environments, showed low responses to eutrophication in the area. Although the mere occurrence of perennial algae did not show clear indicator potential, a distinct discrepancy in the occurrence of bladderwrack, Fucus vesiculosus, was found between two areas with differing eutrophication history, the Bothnian Sea and the Archipelago Sea. The absence of Fucus from many potential sites in the outer Archipelago Sea is likely due to its inability to recover from its disappearance from the area 30-40 years ago, highlighting the importance of past events in macroalgal occurrence. The methodology presented for mapping the potential distribution and the ecological value of reefs showed, that relatively high accuracy in mapping can be achieved by combining existing available data, and the maps produced serve as valuable background information for more detailed surveys. Taken together, the results of the theses contribute significantly to the knowledge on macroalgal communities of the northern Baltic Sea that can be directly applied in various management contexts.

Bacterial community structure and petroleum hydrocarbon degradation in the Baltic Sea

The Baltic Sea is unique by its biological, geochemical and physical features. The number of species of larger organisms is small and the species composition is distinctive. On the contrary microbial communities are diverse. Because of the low salinity levels, bacterial communities differ from the ones in the oceans. Knowing the structure of these communities better and how they response to different environmental conditions helps us to estimate how different factors affect the balance and function of the Baltic Sea ecosystem. Bacteria are the key players when it comes to natural biogeochemical processes and human-induced phenomena like eutrophication, oil spills or disposal of other harmful substances to the sea ecosystem. In this thesis, bacterial community structure in the sea surface microlayer and subsurface water of the Archipelago Sea were compared. In addition, the effect of diatom derived polyunsaturated aldehydes on bacterial community structure was studied by a mesocosm experiment. Diesel, crude oil and polycyclic aromatic hydrocarbon degradation capacity of the Baltic Sea bacteria was studied in smaller scale microcosm experiments. In diesel oil experiments bacteria from water phase of the Archipelago Sea was studied. Sediment and iron manganese concretions collected from the Gulf of Finland were used in the crude oil and polycyclic aromatic hydrocarbon experiments. The amount of polycyclic aromatic hydrocarbon degradation genes was measured in all of the oil degradation experiments. The results show how differences in bacterial community structure can be seen in the sea surface when compared to the subsurface waters. The mesocosm experiment demonstrated how diatom-bacteria interactions depend on other factors than diatom derived polyunsaturated aldehydes, which do not seem to have an effect on the bacterial community structure as has been suggested in earlier studies. The dominant bacterial groups in the diesel microcosms differed in samples taken from a pristine site when compared to a site with previous oil exposure in the Archipelago Sea area. Results of the study with sediment and iron-manganese concretions indicate that there are diverse bacterial communities, typical to each bottom type, inhabiting the bottoms of the Gulf of Finland capable to degrade oil and polycyclic aromatic hydrocarbon compounds.  

Plant-Herbivore Interaction in a Fragmented Landscape: Local Adaptation and Inbreeding

Reciprocal selection between interacting species is a major driver of biodiversity at both the genetic and the species level. This reciprocal selection, or coevolution, has led to the diversification of two highly diverse and abundant groups of organisms, flowering plants and their insect herbivores. In heterogeneous environments, the outcome of coevolved species interactions is influenced by the surrounding community and/or the abiotic environment. The process of adaptation allows species to adapt to their local conditions and to local populations of interacting species. However, adaptation can be disrupted or slowed down by an absence of genetic variation or by increased inbreeding, together with the following inbreeding depression, both of which are common in small and isolated populations that occur in fragmented environments. I studied the interaction between a long-lived plant Vincetoxicum hirundinaria and its specialist herbivore Abrostola asclepiadis in the southwestern archipelago of Finland. I focused on mutual local adaptation of plants and herbivores, which is a demonstration of reciprocal selection between species, a prerequisite for coevolution. I then proceeded to investigate the processes that could potentially hamper local adaptation, or species interaction in general, when the population size is small. I did this by examining how inbreeding of both plants and herbivores affects traits that are important for interaction, as well as among-population variation in the effects of inbreeding. In addition to bi-parental inbreeding, in plants inbreeding can arise from self-fertilization which has important implications for mating system evolution. I found that local adaptation of the plant to its herbivores varied among populations. Local adaptation of the herbivore varied among populations and years, being weaker in populations that were most connected. Inbreeding caused inbreeding depression in both plants and herbivores. In some populations inbreeding depression in herbivore biomass was stronger in herbivores feeding on inbred plants than in those feeding on outbred ones. For plants it was the other way around: inbreeding depression in anti-herbivore resistance decreased when the herbivores were inbred. Underlying some of the among-population variation in the effects of inbreeding is variation in plant phenolic compounds. However, variation in the modification of phenolic compounds in the digestive tract of the herbivore did not explain the inbreeding depression in herbivore biomass. Finally, adult herbivores had a preference for outbred host plants for egg deposition, and herbivore inbreeding had a positive effect on egg survival when the eggs were exposed to predators and parasitoids. These results suggest that plants and herbivores indeed exert reciprocal selection, as demonstrated by the significant local adaptation of V. hirundinaria and A. asclepiadis to one another. The most significant cause of disruption of the local adaptation of herbivore populations was population connectivity, and thus probably gene flow. In plants local adaptation tended to increase with increasing genetic variation. Whether or not inbreeding depression occurred varied according to the life-history stage of the herbivore and/or the plant trait in question. In addition, the effects of inbreeding strongly depended on the population. Taken together, inbreeding modified plant-herbivore interactions at several different levels, and can thus affect the strength of reciprocal selection between species. Thus inbreeding has the potential to affect the outcome of coevolution.

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.

Spatiotemporal Features of Coastal Waters in Southwest Finland

In coastal waters, physico-chemical and biological properties and constituents vary at different time scales. In the study area of this thesis, within the Archipelago Sea in the northern Baltic Sea, seasonal cycles of light and temperature set preconditions for intra-annual variations, but developments at other temporal scales occur as well. Weather-induced runoffs and currents may alter water properties over the short term, and the consequences over time of eutrophication and global changes are to a degree unpredictable. The dynamic characteristics of northern Baltic Sea waters are further diversified at the archipelago coasts. Water properties may differ in adjacent basins, which are separated by island and underwater thresholds limiting water exchange, making the area not only a mosaic of islands but also one of water masses. Long-term monitoring and in situ observations provide an essential data reserve for coastal management and research. Since the seasonal amplitudes of water properties are so high, inter-annual comparisons of water-quality variables have to be based on observations sampled at the same time each year. In this thesis I compare areas by their temporal characteristics, using both inter-annual and seasonal data. After comparing spatial differences in seasonal cycles, I conclude that spatial comparisons and temporal generalizations have to be made with caution. In classifying areas by the state of their waters, the results may be biased even if the sampling is annually simultaneous, since the dynamics of water properties may vary according to the area. The most comprehensive view of the spatiotemporal dynamics of water properties would be achieved by means of comparisons with data consisting of multiple annual samples. For practical reasons, this cannot be achieved with conventional in situ sampling. A holistic understanding of the spatiotemporal features of the water properties of the Archipelago Sea will have to be based on the application of multiple methods, complementing each other’s spatial and temporal coverage. The integration of multi-source observational data and time-series analysis may be methodologically challenging, but it will yield new information as to the spatiotemporal regime of the Archipelago Sea.

Metadatan asialla muistiorganisaatioiden parhaaksi

DC-2014: Metadata Intersections: Bridging the Archipelago of Cultural Memory. Austin, Texas 8.-11.10. http://dcevents.dublincore.org/IntConf/dc-2014.