Abstract This thesis operates at two levels of ecological research, describing the effects of withdrawal and re-introduction of management on grasslands. The aim of the community-level survey was to explore the effects of abandonment, mowing and grazing on semi-natural meadows in northern Finland. At the species level, the aim was to evaluate the responses of three monocarpic meadow species to various degrees of simulated grazing under natural growth conditions. The community-level studies suggest that strongly competitive grass species with rapid vegetative growth, especially those forming tussocks, are able to retain or increase their cover in abandoned meadows. However, most species are able to persist in a meadow for a long time after abandonment, even when a group of immigrants arrive. This leads to a temporary increase in species diversity, and it may therefore be used as an indicator of ongoing succession. Nevertheless, abandonment is harmful for the rare archaeophytic species in the long run. Late mowing does not have extensive short-term impacts on grass-dominated semi-natural meadows. Therefore, it is neither an efficient nor a substitutional way of management when the goal is to restore a formerly grazed pasture. Mowing executed early in the season may, however, be a more appropriate way of inducing changes in species composition and enhancing species richness. According to the results of the species-level studies, Erysimum strictum and Rhinanthus minor tolerate well minor apical damage, while more severe damage has a detrimental impact on the performance of both species. The observed differences in regrowth responses between the two species are presumably due to their different habitat requirements in relation to competition. The species-level experiment with two late-flowering populations of field gentian Gentianella campestris ssp. campestris revealed that the southern, Swedish population that has been regularly grazed and mown overcompensated for the intermediate (50%) damage level, whereas the northern, Finnish field gentians growing in unmanaged habitats showed at best partial or full compensation. Regular grazing and mowing have presumably favoured grazing-tolerant plant species, i.e. species with a good regrowth capacity. Herbivory reshapes grassland plant communities in two ways: directly by affecting the survival and reproductive success of individual plants and indirectly by changing the competitive environment. Tall and competitive perennial species suffer relatively more from damage than true grassland species, i.e. small herbs and grasses, which are better able to tolerate regular tissue losses and respond to damage within the ongoing growing season. As a result, certain species benefit from grazing and mowing in the sense that they may gain more through competitive relaxation than they lose in defoliation.

Abstract Climate change and its warming effects on vegetation and soils are a widely recognized phenomenon. In addition to warming, the understorey vegetation in northern environments has been subjected to several environmental changes, such as increasing nitrogen (N) and other disturbances. This thesis examines the effects of N-fertilization and disturbances on the vegetation biomass and abundance, plant community composition and plant, soil and microbial N and C pools. Seedling establishment of the most common dwarf shrubs (deciduous Vaccinium myrtillus, evergreens V. vitis-idaea and Empetrum nigrum ssp. hermaphroditum) was investigated after artificial disturbance treatments (vegetation and soil removal). These studies were conducted in the boreal and subarctic ecosystems and in the forest-tundra ecotone in northern Finland. N-fertilization and disturbances enhanced the amount of graminoids in plant communities, and the recovery ability of graminoids was enhanced after N-fertilization, which homogenized the vegetation and resulted in a new stable state in the plant community. The recovery ability of evergreen dwarf shrubs was low after disturbances. Disturbances created habitats for seed germination, but the seedling establishment of dwarf shrubs studied was still limited by seed availability. N-fertilization had no effect on microbial biomass. Instead, microbial biomass decreased with disturbance treatment in the boreal forest. However, the concentration of N increased in above-ground vegetation, both after N-fertilization and disturbance without any indication of N immobilization, suggesting that plant species captured the available N effectively for their recovery. The study shows that the likely outcome of N enrichment, when combined with disturbances, is the enhanced growth of graminoids. The seedling establishment does not compensate for the reduction of the vegetative recovery of evergreen dwarf shrubs, which makes evergreen dwarf shrubs sensitive to environmental changes. As the understorey is more resilient to perturbations in the boreal forest than in the subarctic ecosystem, these results emphasize the sensitivity of the vegetation to simultaneous environmental changes in the northernmost ecosystems. Moreover, microbial properties are more resilient to environmental changes than is above-ground vegetation. / Tiivistelmä Ilmaston muutos ja siitä aiheutuvan lämpenemisen vaikutus kasvillisuuteen ja maaperään on laajasti tunnustettu ilmiö. Lämpenemisen lisäksi pohjoisten alueiden aluskasvillisuuteen kohdistuu useita muutospaineita, kuten lisääntynyt typpipitoisuuden nousu ja kasvillisuutta muokkaavat häiriöt. Tässä tutkimuksessa mitattiin lisääntyneen typpipitoisuuden ja häiriöiden vaikutus kasvillisuuden biomassaan ja runsauteen sekä yhteisörakenteeseen, sekä kasvilajeihin, maaperään ja mikrobibiomassaan sitoutuneen typen ja hiilen määrään. Lisäksi tutkittiin yleisimpien varpukasvien (lehtensä pudottava mustikka, ikivihreät puolukka ja variksenmarja) siemenellistä lisääntymistä kokeellisen häiriön (kasvillisuuden tai maaperän poisto) jälkeen. Tutkimukset tehtiin boreaalisessa ja subarctisessa ekosysteemeissä sekä metsänrajaympäristössä Pohjois-Suomessa. Typpilannoitus ja häiriöt lisäsivät heinien määrää kasviyhteisöissä. Lisäksi typpilannoitus edisti heinien kasvullista palautumiskykyä häiriön jälkeen, joka johti kasvillisuuden homogenisoitumiseen ja kasviyhteisön uuteen tasapainotilaan. Häiriöt heikensivät ikivihreiden varpujen kasvullista palautumista häiriön jälkeen. Häiriö loi sopivia elinympäristöjä siementen itämiselle, mutta tutkittujen lajien siementen määrä rajoitti siemenellistä lisääntymistä. Typpilannoitus ei vaikuttanut mikrobibiomassaan, mutta häiriö vähensi mikrobibiomassaa boreaalisessa ekosysteemissä. Kuitenkin kasvien typen pitoisuudet lisääntyivät sekä lannoituksen että häiriön jälkeen ilman viitteitä typen sitoutumisesta mikrobibiomassaan. Tämä viittaa siihen, että kasvit käyttävät maaperän typen tehokkaasti häiriön jälkeiseen palautumiseen. Väitöskirjan mukaan typen lisääntyminen häiriöiden yhteydessä edistää heinien esiintymistä. Koska ikivihreiden varpujen siemenellinen lisääntyminen ei kompensoi häiriöstä kasvulliselle palautumiselle aiheutuvaa haittaa, ovat ikivihreät varvut erityisen herkkiä häiriöille. Aluskasvillisuus on vastustuskykyisempi ympäristön muutoksille boreaalisessa kuin subarktisessa ekosysteemissä, mikä korostaa pohjoisimpien alueiden herkkyyttä yhtäaikaisille ympäristön muutoksille. Maaperän olosuhteet ovat kasvillisuutta kestävämpiä ympäristön muutoksille.