Questions

We aim to answer three questions: (a) what are the effects of canopy shading of different tree species on seed germination of eight understorey grassland species? (b) how is plant species’ occurrence in understorey communities affected by differences in canopy cover and does that depend on tree species composition? and (c) are there mechanistic links between the effects of trees on species’ germination and their occurrence in the understorey plant community?

Location

Semi‐natural wooded pastures in the biosphere reserve Östra Vätterbranterna, southern Sweden.

Methods

In this study, we examined the germination of eight grassland plant species in a seed sowing experiment under natural conditions in the field. Seeds were sown beneath and outside the canopy of two tree species within 48 plots split up by four wooded pasture sites. We combined observed germination responses with a plant community survey to assess the effects of canopy cover in relation to tree species composition on plant community responses. We analysed these data in relation to species’ seed mass and vegetative shade tolerance.

Results

Shade‐tolerant species germinated better beneath compared to outside tree canopies, without any clear advantage of large‐seeded over small‐seeded species. As expected, species’ shade tolerance was also positively related to canopy cover within the understorey plant community. Importantly, we found strong tree species‐specific effects of canopy shading on the species’ germination response, but not on their presence within the plant community. However, optimal canopy cover conditions for germination and for the mature plants differed across grassland species and depended on tree species.

Conclusions

Our results show that different tree species play key ecosystem engineering roles in shaping wooded grassland plant community composition at the germination stage. Management practices favouring specific tree species may therefore be highly relevant for targeted biodiversity conservation of wooded semi‐natural grasslands.

Background: With appropriate management, based on vegetation removal that reverses late-successional vegetation stages, roadsides can support high levels of biodiversity. However, current recommendations for roadside management to conserve or restore biodiversity are largely based on research on non-roadside grassland habitats, and much of the evidence on how roadside management practices affect biodiversity is found in non-peer-reviewed grey literature. Therefore, based on suggestions from key stakeholders and an initial systematic map that identified 97 studies on how biodiversity is affected by vegetation removal on roadsides, we conducted a full systematic review of the effects on plant and invertebrate diversity of disturbance-based maintenance of roadsides. Methods: The review was restricted to effects of non-chemical interventions such as mowing, burning, grazing and mechanical shrub removal. Studies were selected from the systematic map and from an updated search for more recent literature using a priori eligibility criteria. Relevant articles were subject to critical appraisal of clarity and susceptibility to bias, and studies with low or unclear validity were excluded from the review. Data on species richness, species diversity and abundance of functional groups were extracted together with metadata on site properties and other potential effect modifiers. Results from the 54 included studies were summarised in a narrative synthesis, and impacts of mowing practices on the total species richness and diversity of plants and on the abundance of forbs, graminoids and woody plants were quantitatively analysed using t tests of study-level effect ratios. Results: Nearly all of the 54 studies included in the review were conducted in Europe (29) or North America (24). The vast majority of studies (48) examined impacts of mowing. Effects on vascular plants were reported in 51 studies, whereas 8 of the studies reported on invertebrates. Quantitative analysis of plant species richness and species diversity showed that mowing effects were dependent on the interplay between mowing frequency and hay removal. Thus, there were no statistically significant overall effects of mowing vs. no mowing, frequency of mowing, timing of mowing or hay removal. However, species richness was higher in roadsides mowed once or twice per year with hay removal than in unmown roadsides, and positively affected by mowing twice compared to once per year. Similar, but less pronounced, effects were found for plant species diversity. In addition, mowing had a negative impact on woody plant abundance, and increased mowing frequency had a negative impact on graminoid abundance. The few studies on invertebrates showed effects that diverged across taxon groups, and there was not enough data for quantitative analysis of these results. Conclusions: The results provide evidence on the effects of mowing on plant species richness. To increase plant species richness, roadsides should be mowed each year, preferably twice per year, and hay should be removed after each cutting. The review also identifies large knowledge gaps concerning roadside management and its effects on biodiversity, especially regarding invertebrates. Hence, this systematic review provides not only a valuable basis for evidence-based management but also guidance for future research on this topic, essential to inform management of road networks for biodiversity conservation.

Europe's woodland and savanna rangelands, often part of silvopastoral systems known as wood-pastures, are deteriorating because of abandonment that leads to return to a forested state or lack of tree regeneration from overgrazing or tree and shrub removal. Despite numerous local studies, there has been no broader survey of the stand structure of European wood-pastures showing which systems are at risk of losing their semiopen character. This overview aims to 1) show some of the differences and similarities in wood-pastures from landscapes across Europe and 2) identify which of these wood-pastures are at risk of losing their semiopen character. We collated a dataset of 13 693 trees from 390 plots in wood-pastures from eight different European regions (western Estonia, eastern Greece, northern Germany, Hungary, northern Italy, southern Portugal, central Romania, and southern Sweden), including tree diameters at breast height, tree density, management type, and tree species composition. On the basis of their structural characteristics, we classified wood-pastures using principal component analysis (PCA) and cluster analysis. The PCA showed a gradient from dense wood-pastures with high levels of regeneration (e.g., in Estonia) to sparse wood-pastures with large trees but a lack of regeneration (e.g., in Romania). Along this gradient, we identified three main groups of wood-pastures: 1) sparse wood-pastures with mostly big trees; 2) dense wood-pastures composed of small trees, and 3) wood-pastures containing a wide range of tree ages. Our results show a large structural gradient in European wood-pastures, as well as regeneration problems varying in their severity, highlighting the importance of social-ecological context for wood-pasture conditions. To maintain the ecological and cultural integrity of European wood-pastures, we suggest 1) more comprehensively considering them in European policies such as the Common Agricultural Policy and EU Habitats Directive, while 2) taking into account their structural characteristics and social-ecological backgrounds.

Wood pastures are some of the most species-rich environments found in Europe and therefore essential habitats for biodiversity conservation. Society also puts faith in multiple values of trees, ranging from climate change mitigation to socio-cultural traditions. Therefore, the seemingly arbitrary tree density limit for pasture environments imposed by the EU through its Common Agricultural Policy (CAP) threatened both biological and societal values. In this study on farmers' perspectives, we target the effects of the CAP tree density limit on management of wood pastures in a low-intensively managed agricultural landscape of southern Sweden. The case of simplifying nature by using simple number limitations was used as an entry point in semi-structured, open-ended, interviews with farmers and officials about their view on trees and pasture management in relation to policy directives. The interviews showed that the policy incentive shifted the management focus from grazing quality to the number of trees and that farmers are willing to cut in order to get subsidies and timber revenues, however not unreflectingly. Farmers had high knowledge about the wide ranging social, cultural and natural values of trees, and are often themselves as good regulators of tree management as policies intend to be. Our study reveals many difficulties in managing the complex relations within landscapes with simplified legal measures, opening up for further discussion about improving policy instruments to preserve both social and biological values of wood pastures. However, although the tree density limit has been criticised on many points related to biodiversity conservation, this study shows that other values linked to pasture trees, e.g. the aesthetic values and their importance as shelter for grazing animals, could be an argument to actually keep the focus on trees as indicators of pasture management quality. We suggest that trees in general and wood-pastures in particular therefore are good starting points, or boundary-objects, for collaboration between production and conservation interests of farming and environmental management.

Worldwide, biodiversity conservation is one of the key challenges for a sustainable future of nature and society. It is particularly important to preserve high quality habitats within otherwise intensively managed agricultural landscapes. Within the European Union (EU), farmers are highly dependent on agricultural subsidies through the Common Agricultural Policy (CAP), which hence have a strong influence on management and biodiversity. In European agricultural landscapes, wooded pastures form important habitats that contribute to landscape level heterogeneity and high local biodiversity, values which are often closely linked to trees. Unfortunately, many of these values were put at risk when a tree density limitation was introduced within the CAP, encouraging farmers to keep pastures open and ensuring grazing management. However, limiting tree density to a specific number to increase biodiversity finds little basis in the scientific literature. The main objective of this thesis is therefore to investigate how different measures of biodiversity across multiple taxa are affected by tree density and to study the farmers' perspective on this CAP regulation. Wooded pastures in the biosphere reserve Östra Vätterbranterna in southern Sweden were used as study sites. This thesis shows that encouraging farmers to cut trees to receive subsidies weakens the link between social and ecological values of wooded pastures, with potential subsequent losses in biodiversity. Trees were almost exclusively positive for biodiversity within this study system, increasing the species richness of plants, birds and bats. However, functional diversity across these taxa were mainly affected by other vegetation attributes within and around the pastures, such as shrub density and surrounding forest cover. A seed sowing experiment showed how trees partly shape plant communities already at the germination stage. Further, responses of functional diversity was mainly driven by resource use related traits among plants and birds, whereas bat functional diversity responses were mainly determined by their ability to manoeuvre through the micro-habitats of wooded pastures. Based on this thesis, I conclude that the tree density limit proposed by the EU has failed to capture the unique biological values of continuously managed wooded pastures and that the social-ecological links between policy, management and biodiversity are threatened by number specific governance of nature. It is therefore promising that the EU in November 2017 announced to open up for excluding the tree density focus in the CAP. Further development of the CAP can benefit from the findings of this thesis, revealing important knowledge gaps on biodiversity patterns in relation to trees in pastures.

The European Union's Common Agricultural Policy (CAP) recommends subsidies are only granted for wood-pastures with < 100 trees/ha. This guidance exists despite these habitats being among the most biodiverse in boreal Europe and currently under threat due to land conversion. Bats are important bio-indicators of agricultural landscapes, but bat diversity has not explicitly been studied in relation to this policy. We investigate how bat activity, foraging, species richness and functional groups are affected in twenty-six wood-pastures along a gradient of tree density, from open to dense. In parallel, open fields and deciduous forests were sampled and the effect of the surrounding landscape configuration was explored. Our results show a consistent increase in total bat activity, foraging activity and species richness within wood-pastures along the tree density gradient. We find optimal tree densities within wood-pastures are higher than values reported in previous studies, and suggest thresholds might depend on the landscape context. Shrub density was a strong predictor of total bat activity and foraging; whilst structural variation of tree size in wood-pastures was most strongly correlated with species richness. We show that wood-pastures are an important habitat and in comparison to forests they contribute to higher bat species richness and activity levels. Interestingly, higher activity levels of forest feeding specialists were observed in wood-pastures compared to forests. At the landscape level, amount of water in the landscape was the strongest predictor of bat activity whilst deciduous forest mostly influenced foraging activity. This study demonstrates that tree density within wood-pastures is not a limiting factor of bat activity and foraging and that other habitat and landscape parameters are important. Thereby focusing solely on tree density limits will not help to promote the ecological requirements for bats. Instead we suggest that a results based approach to CAP payments would be preferable.

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.

Background: In many parts of the world, roadsides are regularly managed for traffic-safety reasons. Hence, there are similarities between roadsides and certain other managed habitats, such as wooded pastures and mown or grazed grasslands. These habitats have declined rapidly in Europe during the last century. For many species historically associated with them, roadsides may function as new primary habitats or as dispersal corridors in fragmented landscapes. Current recommendations for roadside management to promote conservation values are largely based on studies of plants in semi-natural grasslands, although such areas often differ from roadsides in terms of environmental conditions and disturbance regimes. Moreover, roadsides provide habitat not only for plants but also for many insects. For these reasons, stakeholders in Sweden have emphasised the need for more targeted management recommendations, based on actual studies of roadside biodiversity. Methods: This systematic map provides an overview of the available evidence on how biodiversity is affected by various forms of roadside management, and how such management influences the dispersal of species along roads or roadsides. We searched for literature using 13 online publication databases, 4 search engines, 36 specialist websites and 5 literature reviews. Search terms were developed in English, Danish, Dutch, French, German, Norwegian, Spanish and Swedish. Identified articles were screened for relevance using criteria set out in a protocol. No geographical restrictions were applied, and all species and groups of organisms were considered. Descriptions of included studies are available in an Excel file, and also in an interactive GIS application that can be accessed at an external website. Results: Our searches identified more than 7000 articles. The 207 articles included after screening described 301 individual studies considered to be relevant. More than two-thirds of these studies were conducted in North America, with most of the rest performed in Europe. More than half of the studies were published in grey literature such as reports from agencies or consultants. The interventions most commonly studied were herbicide use, sowing and mowing, followed by soil amendments such as mulching and fertiliser additions. The outcomes most frequently reported were effects of interventions on the abundance or species richness of herbs/forbs, graminoids and woody plants. Effects on insects and birds were reported in 6 and 3% of the studies, respectively. Conclusions: This systematic map is based on a comprehensive and systematic screening of all available literature on the effects of roadside management on biodiversity and dispersal of species. As such it should be of value to a range of actors, including managers and policymakers. The map provides a key to finding concrete guidance for conservation- and restoration-oriented roadside management from published research. However, the map also highlights important knowledge gaps: little data was found for some geographical regions, research is heavily biased taxonomically towards plants, and no study was found on how species dispersal was affected by roadside management. The map could therefore be a source of inspiration for new research.

Background: Roadsides have been acknowledged as potential substitutes for semi-natural grasslands and other open habitats with high biodiversity, many of which are now declining. Current recommendations for roadside management to promote conservation of biodiversity are largely based on studies of plants in meadows or pastures, although such areas often differ from roadsides in terms of environmental conditions and disturbance regimes. Stakeholders in Sweden have emphasised the need for more targeted guidelines for roadside management, based on actual roadside studies. We recently performed a systematic mapping of the evidence on how roadside management affects biodiversity and the dispersal of species. Through this process, we found 98 studies on how the richness or abundance of species on roadsides is affected by management such as regular mowing, burning, grazing or selective mechanical removal of plants. Since all of these interventions entail removal of plant biomass, they are comparable. Most of the studies recorded management effects on vascular plants, but there were 14 investigations of insects and other invertebrates. We now intend to proceed with a full systematic review of how maintenance or restoration of roadsides based on non-chemical vegetation removal affects the diversity of vascular plants and invertebrates. Methods: Most of the evidence on which the proposed systematic review is to be based will be selected from the systematic map. To identify more recently published literature on the topic of the review, we will perform a search update using a subset of the search terms applied for the systematic map. The criteria for inclusion of studies will be the same as for the map, except that interventions and outcomes will be restricted to those relevant to the review. Relevant studies will be subject to critical appraisal and categorised as having high or low validity for the review. Studies with low validity will be excluded from the review. Utilisable data on outcomes, interventions and other potential effect modifiers will be extracted from included articles. A narrative synthesis will describe the validity and findings of all studies in the review. Where a sufficient number of studies report similar outcome types, meta-analysis will be conducted.

1. Recent reforms in the Common Agricultural Policy aim for a greening of the subsidy system with potential improvements for biodiversity conservation. As part of that process, the tree density limit for pastures to qualify for European Union subsidies has been increased from 50 to 100 trees per hectare. However, recent studies show that the high biodiversity values of these habitats may be threatened by these limits, highlighting the need for policy improvements. Still, little is known about the direct effects of tree density limitations on bird communities in woody pastures. 2. We investigated how bird diversity and species composition are affected by tree density in 49 Swedish woody pastures along a gradient of 4-214 trees per hectare. We recorded bird communities, tree density and stand structure estimates in the field and estimated forest cover in the surrounding landscape from aerial photos. Using generalised additive models and redundancy analysis, we analysed how bird territorial species richness, bird abundance and species composition are affected by tree density, taking into account other local and landscape scale explanatory variables. 3. Tree density had a significant positive effect on bird species richness at low tree densities and species richness saturated at approximately 50 trees per hectare. Shrub density had a significant positive linear effect on both bird species richness and abundance. Tree and shrub density were also the major drivers of bird community composition, with secondary effects of tree species diversity and landscape forest cover. 4. Policy implications. Our results show that tree density is not the limiting factor, but rather a driver of bird diversity and species composition in woody pastures and that tree density limits may fail to capture the whole range of biological values. To improve policy recommendations, we therefore stress the importance of considering additional social-ecological drivers associated to management quality, e.g. taking into account moral and cultural motivations among farmers, to preserve biodiversity in woody pastures.