Biodiversity loss and its effects on humanity is of major global concern. While a growing body of literature confirms positive relationships between biodiversity and multiple ecological functions, the links between biodiversity, ecological functions and multiple ecosystem services is yet unclear. Studies of biodiversity-functionality relationships are mainly based on computer simulations or controlled field experiments using only few species. Here, we use a trait-based approach to integrate plant functions into an ecosystem service assessment to address impacts of restoration on species-rich grasslands over time. We found trade-offs among functions and services when analysing contributions from individual species. At the community level, these trade-offs disappeared for almost all services with time since restoration as an effect of increased species diversity and more evenly distributed species. Restoration to enhance biodiversity also in species-rich communities is therefore essential to secure higher functional redundancy towards disturbances and sustainable provision of multiple ecosystem services over time.

In line with the 2010 Aichi Convention for Biological Diversity, the European Union has a goal to restore 15% of degraded ecosystems and their services by the year 2020 (target 2, Europe 2020). This includes restoration of semi-natural grasslands (SNGs). Management of both intact and restored SNGs is dependent on people's willingness to manage them. Due to low profitability, management abandonment still occurs all over Europe, which highlights the need to raise farmers' and landowners' perspectives. In this study, we combined survey data and in-depth interviews with farmers/landowners managing previously restored SNGs, to understand how they perceive the restoration process, the outcome and future management. Survey and interview data were analysed in relation to biodiversity and Agri-environmental payments data from the restored sites. Almost all respondents considered the restoration successful and the re-inventoried restored SNGs also showed an increase in plant diversity. Nevertheless, 10% of the restored SNGs were abandoned again post-restoration and 40% of the respondents were unsure if they would continue the management in the future. Abandoned management may cause a negative trend in terms of decreased biological, cultural and aesthetic values, in the local community, as well as for the society in general. Most respondents explained a strong dependency on Agri-environmental payments, both as a restoration incentive and for post-restoration management. Also non-financial support from authorities in form of feedback and advice was requested, as well as support from the local community and society as a whole. Future management in a longer time perspective was strongly coupled to the farm economy, i.e. received Agri-environmental payments at farm-level and profit from selling agricultural products, and whether the farmers had successors. We conclude that both social and ecological factors, here farm economy, authority support and proper management, must be in place for long-term success of grassland restoration.

Humans play a major role shaping the living conditions for not only ourselves, but also all other species on Earth. In fact, some species-rich habitat types require human management to uphold the biodiversity and related ecosystem services. One of the world’s most biodiverse habitats on small spatial scales, semi-natural grasslands, have been formed over the course of centuries through extensive grazing and mowing. However, due to political and economic reasons, up to 90% of the European semi-natural grasslands have been lost during the 20th century. To counteract these drastic losses, restoration actions are implemented in environmental policies across Europe. Yet, knowledge of the long-term restoration effects on biodiversity and ecosystem services is still limited. The vast need for future restoration also requires a better understanding of how different pre-conditions affect the restoration outcome, as well as how stakeholders perceive restoration, to be able to prioritise between sites and recognise the limitations of the restoration process. In this thesis, I examine restoration outcomes in Swedish semi-natural grasslands, in terms of plant diversity, associated ecosystem services and from the farmers’ and land-owners’ perspective. The outcome is also analysed in relation to environmental factors at the local and landscape scale. I found that the overall community composition recovered to resemble intact reference communities, but it took relatively long time (12-20 years). Moreover, the reference sites still had higher species richness both at large and small spatial scales, more grassland specialist species and a higher abundance of plant species important to the five tested ecosystem services (meat production, pollination, water retention, temperature regulation and cultural heritage). My results show that prioritising large, unfertilised, newly abandoned grasslands situated in landscapes containing a large grassland specialist species pool and high amounts of intact and remnant semi-natural grasslands, could speed up the plant recovery. However, prioritising fast results does not necessarily ensure long-term success at a larger spatial scale. Since restoration success can be interpreted differently depending on evaluation measure used, pre-defined, clear and realistic goals are essential. While the surveyed farmers and landowners overall perceived the restoration as successful, 40% were unsure whether the grasslands will be managed in the future. Low profitability still poses a threat to their maintenance and thus, also to the coupled biodiversity and ecosystem services. Policy changes are therefore urgently needed to facilitate incentives for sustained management of restored and intact European semi-natural grasslands in a long-term perspective.

Människan har sedan länge påverkat jordens ekosystem, där vissa habitat numer är beroende av skötsel för att upprätthålla biologisk mångfald och ekosystemtjänster. I naturbetesmarker och ängar (traditionella gräsmarker) har århundranden av hävd genom bete och slåtter format en artrikedom att jämföra med tropisk regnskog, med skillnaden att många av dessa arter är beroende av hävden för sin överlevnad. På grund av det senaste århundrandets jordbruksintensifiering har dock upp till 90% av de europeiska gräsmarkerna vuxit igen eller omvandlats till åkermark eller skogsplantering. För att motverka detta är restaurering av dessa habitat en viktig åtgärd som ingår i bl.a. EU:s ekonomiska stödsystem till lantbrukare. Ekonomiska incitament för restaurering har funnits i flera decennier, men kunskapen om långsiktiga effekter på biologisk mångfald och ekosystemtjänster är fortfarande relativt begränsad. För att möta det omfattande framtida restaureringsbehovet behövs också mer information om hur olika miljöfaktorer kan påverka resultatet, samt hur berörda aktörer uppfattar restaureringen. I denna avhandling undersöker jag restaurering av svenska traditionellt hävdade gräsmarker, med fokus på mångfald av växter och ekosystemtjänster, samt ur lantbrukares och markägares perspektiv. Detta analyseras i förhållande till lokala förhållanden i gräsmarkerna och omgivande landskapsfaktorer. Jag fann att artsammansättningen i restaurerade gräsmarker utvecklades till att likna sammansättningen i kontinuerligt hävdade (intakta) referensgräsmarker, men att denna återhämtning tog relativt lång tid (12-20 år). Referensmarkerna hade dock fortfarande högre artrikedom, fler gräsmarksspecialiserade arter och högre abundans av växter viktiga för fem ekosystemtjänster: köttproduktion, pollinering, vattenhållningsförmåga, kolinlagring och kulturarv. Mina resultat visade även att prioritering av stora, ogödslade, nyligen övergivna gräsmarker belägna i landskap med många intakta gräsmarker och resthabitat, samt en stor artpool med gräsmarksspecialister, skulle kunna påskynda växters återetablering. Det bör dock påpekas att restaurering av gräsmarker kan ge goda effekter också i andra slags landskap, även om det kan ta längre tid. Sammantaget synliggör min avhandling att beroende på vilken utvärderingsmetod som används kan restaureringsresultatet tolkas på olika sätt. För att kunna göra en korrekt utvärdering behövs därför tydliga och realistiska mål som definierats innan restaureringen. Lantbrukarna och markägarna upplevde överlag att restaureringen varit framgångsrik. Trots detta var 40% osäkra på huruvida dessa gräsmarker kommer att skötas i framtiden, framför allt på grund av låg lönsamhet. Därmed är det av yttersta vikt att stödsystemen utformas för att gynna hållbar och långsiktig hävd av både restaurerade och intakta naturbetesmarker och ängar i Europa.

Habitat loss through land-use change is the most pressing threat to biodiversity worldwide. European semi-natural grasslands have suffered an ongoing decline since the early twentieth century, but we have limited knowledge of how grassland loss has affected biodiversity across large spatial scales. We quantify land-use change over 50-70 years across a 175,000 km(2) super-region in southern Sweden, identifying a widespread loss of open cover and a homogenisation of landscape structure, although these patterns vary considerably depending on the historical composition of the landscape. Analysing species inventories from 46,796 semi-natural grasslands, our results indicate that habitat loss and degradation have resulted in a decline in grassland specialist plant species. Local factors are the best predictors of specialist richness, but the historical landscape predicts present-day richness better than the contemporary landscape. This supports the widespread existence of time-lagged biodiversity responses, indicating that further species losses could occur in the future.

Habitat restoration is an important complement to protecting habitat for the conservation of biodiversity. Semi-natural grasslands are target habitats for ecological restoration in temperate Europe. Restoration of abandoned semi-natural grasslands often relies on spontaneous colonisation of plant species from the soil seed bank or the surrounding landscape. Although many studies show that the regional species pool is important for upholding local diversity, its effect on restoration outcome in semi-natural grasslands is poorly known. In this multi landscape study, we examined grassland specialist species occurring in restored grasslands and the effect of specialist species pool, landscape composition and local temporal factors. We found that specialist richness and frequency was positively affected by specialist richness and frequency in the surrounding landscape. Specialist richness in the restored grasslands also increased with time since restoration. Moreover, specialist frequency in the restored grassland increased with the proportion of semi-natural and remnant grassland habitats in the landscape. We also found a positive relationship between the proportion of species occurring in both the restored grassland and its surrounding landscape and time since restoration, in landscapes with high proportions of semi natural grasslands. This suggests that both temporal factors, as well as the landscape composition and species pool, affect plant recolonisation in restored semi-natural grasslands.

Habitat destruction and degradation represent serious threats to biodiversity, and quantification of land-use change over time is important for understanding the consequences of these changes to organisms and ecosystem service provision. Comparing land use between maps from different time periods allows estimation of the magnitude of habitat change in an area. However, digitizing historical maps manually is time-consuming and analyses of change are usually carried out at small spatial extents or at low resolutions. HistMapR contains a number of functions that can be used to semi-automatically digitize historical land use according to a map's colours, as defined by the RGB bands of the raster image. We test the method on different historical land-use map series and compare results to manual digitizations. Digitization is fast, and agreement with manually digitized maps of around 80-90% meets common targets for image classification. We hope that the ability to quickly classify large areas of historical land use will promote the inclusion of land-use change into analyses of biodiversity, species distributions and ecosystem services.

Restoration is important for biodiversity conservation worldwide, but surprisingly little is known about its efficiency in a long-term perspective. In this study, we re-examined Swedish semi-natural grasslands 12-20 years after the restoration, by comparing field inventories of vascular plant species diversity made in 2001 with follow-up inventories in 2012. We also analysed restoration effect in relation to six environmental factors and used continuously managed semi-natural grasslands as references of desired state after restoration. We found that total species richness increased over time but not to reference levels, while there were no significant changes in species density or number of grassland specialists. However, the overall species composition in the restored sites, as well as grassland specialist composition, now largely resembled reference conditions. Fertilisation and time between abandonment and restoration were the only environmental variables that affected total species composition change, while site area affected change in grassland specialist composition. Our results show that restoration of semi-natural grasslands can contribute to conservation of semi-natural habitats and their associated biodiversity. Yet, due to the vague restoration goals for these sites, it is difficult to evaluate the restoration success, which emphasise the general need for clear and measurable goals.