Short-term plant-decomposer feedbacks in grassland plants

Plant species differ in their effects on ecosystem productivity and it is recognised that these effects are partly due to plant species-specific influences on soil processes. Until recently, however, not much attention was given to the potential role played by soil biota in these species-specific effects. While soil decomposers are responsible for governing the availability of nutrients for plant production, they simultaneously depend on the amount of carbon provided by plants. Litter and rhizodeposition constitute the two basal resources that plants provide to soil decomposer food webs. While it has been shown that both of these can have effects on soil decomposer communities that differ among plant species, the putative significance of these effects for plant nitrogen (N) acquisition is currently understudied. My PhD work aimed at clarifying whether the species-specific influences of three temperate grassland plants on the soil microfood-web, through rhizodeposition and litter, can feed back to plant N uptake. The methods and approach used (15N labelling of plant litter in microcosm experiments) revealed to be an effective combination of tools in studying these feedbacks. Plant effects on soil organisms were shown to differ significantly between plant species and the effects could be followed across several trophic levels. The labelling of litter further permitted the evaluation of plant acquisition of N derived from soil organic matter. The results show that the structure of the soil microfood-web can have a significant role in plant N acquisition when the structure is experimentally manipulated, such as when comparing systems consisting of microbes to those consisting of microbes and their grazers. However, despite this, the results indicate that differences in N uptake from soil organic matter between different plant species are not related to the effects these species exert on the structure of the soil microfood-web. Rather, these differences in N uptake seem to be determined by other species-specific traits of live plants and their litter. My results thus indicate that different resources provided by different plant species may not induce species-specific decomposer feedbacks on plant N uptake from soil organic matter. This further suggests that the species-specific plant effects on soil decomposer communities may not, at least in the short term, have significant consequences on plant production.

Maintaining plant species richness by cattle grazing - mesic semi-natural grasslands as focal habitats

Semi-natural grasslands are the most important agricultural areas for biodiversity. The present study investigates the effects of traditional livestock grazing and mowing on plant species richness, the main emphasis being on cattle grazing in mesic semi-natural grasslands. The two reviews provide a thorough assessment of the multifaceted impacts and importance of grazing and mowing management to plant species richness. It is emphasized that livestock grazing and mowing have partially compensated the suppression of major natural disturbances by humans and mitigated the negative effects of eutrophication. This hypothesis has important consequences for nature conservation: A large proportion of European species originally adapted to natural disturbances may be at present dependent on livestock grazing and / or mowing. Furthermore, grazing and mowing are key management methods to mitigate effects of nutrient-enrichment. The species composition and richness in old (continuously grazed), new (grazing restarting 3-8 years ago) and abandoned (over 10 years) pastures differed consistently across a range of spatial scales, and was intermediate in new pastures compared to old and abandoned pastures. In mesic grasslands most plant species were shown to benefit from cattle grazing. Indicator species of biologically valuable grasslands and rare species were more abundant in grazed than in abandoned grasslands. Steep S-SW-facing slopes are the most suitable sites for many grassland plants and should be prioritized in grassland restoration. The proportion of species trait groups benefiting from grazing was higher in mesic semi-natural grasslands than in dry and wet grasslands. Consequently, species trait responses to grazing and the effectiveness of the natural factors limiting plant growth may be intimately linked High plant species richness of traditionally mowed and grazed areas is explained by numerous factors which operate on different spatial scales. Particularly important for maintaining large scale plant species richness are evolutionary and mitigation factors. Grazing and mowing cause a shift towards the conditions that have occurred during the evolutionary history of European plant species by modifying key ecological factors (nutrients, pH and light). The results of this Dissertation suggest that restoration of semi-natural grasslands by private farmers is potentially a useful method to manage biodiversity in the agricultural landscape. However, the quality of management is commonly improper, particularly due to financial constraints. For enhanced success of restoration, management regulations in the agri-environment scheme need to be defined more explicitly and the scheme should be revised to encourage management of biodiversity.

Dynamics of Finnish starlings in 1951-2005 : from monitoring to population modelling

Farmland bird species have been declining in Europe. Many declines have coincided with general intensification of farming practices. In Finland, replacement of mixed farming, including rotational pastures, with specialized cultivation has been one of the most drastic changes from the 1960s to the 1990s. This kind of habitat deterioration limits the persistence of populations, as has been previously indicated from local populations. Integrated population monitoring, which gathers species-specific information of population size and demography, can be used to assess the response of a population to environment changes also at a large spatial scale. I targeted my analysis at the Finnish starling (Sturnus vulgaris). Starlings are common breeders in farmland habitats, but severe declines of local populations have been reported from Finland in the 1970s and 1980s and later from other parts of Europe. Habitat deterioration (replacement of pasture and grassland habitats with specialized cultivation areas) limits reproductive success of the species. I analysed regional population data in order to exemplify the importance of agricultural change to bird population dynamics. I used nestling ringing and nest-card data from 1951 to 2005 in order to quantify population trends and per capita reproductive success within several geographical regions (south/north and west/east aspects). I used matrix modelling, acknowledging age-specific survival and fecundity parameters and density-dependence, to model population dynamics. Finnish starlings declined by 80% from the end of the 1960s up to the end of the 1980s. The observed patterns and the model indicated that the population decline was due to the decline of the carrying capacity of farmland habitats. The decline was most severe in north Finland where populations largely become extinct. However, habitat deterioration was most severe in the southern breeding areas. The deteriorations in habitat quality decreased reproduction, which finally caused the decline. I suggest that poorly-productive northern populations have been partly maintained by immigration from the highly-productive southern populations. As the southern populations declined, ceasing emigration caused the population extinction in north. This phenomenon was explained with source sink population dynamics, which I structured and verified on the basis of a spatially explicit simulation model. I found that southern Finnish starling population exhibits ten-year cyclic regularity, a phenomenon that can be explained with delayed density-dependence in reproduction.

Management of semi-natural grasslands for butterfly and moth communities

This work focuses on the factors affecting species richness, abundance and species composition of butterflies and moths in Finnish semi-natural grasslands, with a special interest in the effects of grazing management. In addition, an aim was set at evaluating the effectiveness of the support for livestock grazing in semi-natural grasslands, which is included in the Finnish agri-environment scheme. In the first field study, butterfly and moth communities in resumed semi-natural pastures were com-pared to old, annually grazed and abandoned previous pastures. Butterfly and moth species compo-sition in restored pastures resembled the compositions observed in old pastures after circa five years of resumed cattle grazing, but diversity of butterflies and moths in resumed pastures remained at a lower level compared with old pastures. None of the butterfly and moth species typical of old pas-tures had become more abundant in restored pastures compared with abandoned pastures. There-fore, it appears that restoration of butterfly and moth communities inhabiting semi-natural grass-lands requires a longer time that was available for monitoring in this study. In the second study, it was shown that local habitat quality has the largest impact on the occurrence and abundance of butterflies and moths compared to the effects of grassland patch area and connec-tivity of the regional grassland network. This emphasizes the importance of current and historical management of semi-natural grasslands on butterfly and moth communities. A positive effect of habitat connectivity was observed on total abundance of the declining butterflies and moths, sug-gesting that these species have strongest populations in well-connected habitat networks. Highest species richness and peak abundance of most individual species of butterflies and moths were generally observed in taller grassland vegetation compared with vascular plants, suggesting a preference towards less intensive management in insects. These differences between plants and their insect herbivores may be understood in the light of both (1) the higher structural diversity of tall vegetation and (2) weaker tolerance of disturbances by herbivorous insects due to their higher trophic level compared to plants. The ecological requirements of all species and species groups inhabiting semi-natural grasslands are probably never met at single restricted sites. Therefore, regional implementation of management to create differently managed areas is imperative for the conservation of different species and species groups dependent on semi-natural grasslands. With limited resources it might be reasonable to focus much of the management efforts in the densest networks of suitable habitat to minimise the risk of extinction of the declining species.

Defoliation and patchy nutrient return drive grazing effects on plant and soil properties in a dairy cow pasture

Large herbivores can influence plant and soil properties in grassland ecosystems, but especially for belowground biota and processes, the mechanisms that explain these effects are not fully understood. Here, we examine the capability of three grazing mechanisms-plant defoliation, dung and urine return, and physical presence of animals (causing trampling and excreta return in patches)-to explain grazing effects in Phleum pratense-Festuca pratensis dairy cow pasture in Finland. Comparison of control plots and plots grazed by cows showed that grazing maintained original plant-community structure, decreased shoot mass and root N and P concentrations, increased shoot N and P concentrations, and had an inconsistent effect on root mass. Among soil fauna, grazing increased the abundance of fungivorous nematodes and Aporrectodea earthworms and decreased the abundance of detritivorous enchytraeids and Lumbricus earthworms. Grazing also increased soil density and pH but did not affect average soil inorganic-N concentration. To reveal the mechanisms behind these effects, we analyzed results from mowed plots and plots that were both mowed and treated with a dung and urine mixture. This comparison revealed that grazing effects on plant attributes were almost entirely explained by defoliation, with only one partly explained by excreta return. Among belowground attributes, however, the mechanisms were more mixed, with effects explained by defoliation, patchy excreta return, and cow trampling. Average soil inorganic-N concentration was not affected by grazing because it was simultaneously decreased by defoliation and increased by cow presence. Presence of cows created great spatial heterogeneity in soil N availability and abundance of fungivorous nematodes. A greenhouse trial revealed a grazing-induced soil feedback on plant growth, which was explained by patchiness in N availability rather than changes in soil biota. Our results show that grazing effects on plant attributes can be satisfactorily predicted using the effects of defoliation, whereas those on soil fauna and soil N availability need understanding of other mechanisms as well. The results indicate that defoliation-induced changes in plant ecophysiology and the great spatial variation in N availability created by grazers are the two key mechanisms through which large herbivores can control grassland ecosystems.

Luonnonhoitopellot maatalousympäristön luonnon monimuotoisuuden edistämisessä

Luonnonhoitopellot lisättiin uutena vapaaehtoisena toimenpiteenä maatalouden ympäristötukeen vuonna 2009. Luonnonhoitopeltoihin kuuluvat monivuotiset nurmipellot sekä niitty-, riista- ja maisemakasveilla kylvetyt pellot. Toimenpiteen avulla pyritään suojelemaan ja lisäämään maatalousympäristön luonnon monimuotoisuutta sekä muun muassa vähentämään maatalouden ravinnehuuhtoumia. Vuonna 2010 luonnonhoitopeltoja oli yli seitsemän prosenttia Suomen viljelyalasta. Luonnonhoitopeltojen ympäristövaikutusten arvioiminen ja toimenpiteen kehittäminen on tärkeää, jotta toimenpiteeseen käytettävät varat eivät valu hukkaan. Tämän tutkimuksen tavoitteena oli selvittää, millainen merkitys luonnonhoitopelloilla on maatalousympäristön monimuotoisuuden kannalta; millaiset luonnonhoitopellot ovat monimuotoisuuden kannalta arvokkaimpia; ja kuinka toimenpidettä kannattaa kehittää. Kysymyksiin pyrittiin vastaamaan tutkimalla putkilokasvilajistoa, -lajirikkautta ja kasvillisuuden rakennetta erilaisilla luonnonhoitopelloilla, sekä vertaamalla luonnonhoitopeltojen kasvillisuutta pientareiden ja niittyjen kasvillisuuteen. Maastotyö luonnonhoitopelloilla tehtiin kesällä 2010 Uudenmaan ja Pohjois-Pohjanmaan ELY-keskusalueilla. Lisäksi tutkimuslohkoista kerättiin tietoja viljelijäkyselyn kautta. Niittypellot osoittautuivat tutkimuksessa lajirikkaimmaksi luonnonhoitopeltotyypiksi, mikä selittynee lähinnä sillä, että ne kylvetään heikosti kilpailukykyisellä siemenseoksella. Nurmi- ja niittypelloilla lohkon viljavuus korreloi negatiivisesti lajirikkauden kanssa. Lajikoostumukseltaan kaikki luonnonhoitopellot eroavat toisistaan sekä maatalousympäristön puoliluonnontilaisista elinympäristöistä, ja lisäävät siten monimuotoisuutta maisematasolla. Luonnonhoitopeltojen lajisto ei ole kuitenkaan suojelun kannalta erityisen arvokasta. Toimenpiteen maatalousympäristöä rikastuttavaa vaikutusta vähentää, että tällä hetkellä valtaosa luonnonhoitopelloista on monivuotisia nurmipeltoja. Luonnonhoitopeltojen siemenseoksia kehittämällä, perustamis- ja hoitomenetelmiä tutkimalla ja neuvontaan panostamalla voidaan lisätä luonnonhoitopeltojen arvoa niin luonnon, viljelijän kuin yhteiskunnankin kannalta.