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1.Maintaining semi-natural open habitats requires biomass removal, which can be achieved by extensive grazing with livestock species. However, implementing this established conservation management strategy in large or access-restricted areas is often not possible. 2.We investigated grazing by wild and free-ranging red deer Cervus elaphus as an alternative conservation management approach on an active military training area in Germany. In grasslands and heathlands protected under the EU Habitats Directive, we quantified above-ground net primary productivity, forage quality and forage removal by red deer over three successive years. To assess synergistic effects between wildlife grazing and additional grassland management measures removing vegetation biomass (i.e. burning, mowing), we conducted our grazing experiment in burnt, mown and untreated grasslands.3. Annual forage removal by red deer amounted to 35%, 44%, 48% and 59% of the above-ground net primary productivity in burnt, mown and untreated grasslands, and untreated heathlands, respectively. Theoretically, a similar annual biomass removal could be obtained by livestock grazing with 0.54 animal units ha −1 in grasslands and 0.45 animal units ha −1 in heathlands. In grasslands, daily rates of forage removal peaked in spring and early summer, whereas in heathlands, forage removal rates were only significant in winter. 4.Forage removal rates in grasslands increased with productivity and forage quality, which were both enhanced in mown grasslands. This suggests mowing can increase grassland attractiveness to red deer. Forage removal rates in heathlands did not relate to productivity or forage quality but to standing biomass. Synthesis and applications.We demonstrate that the quantity of forage removed by wild red deer can be comparable to that of livestock in common conservation grazing systems. The seasonal patterns of forage removal by red deer in grasslands and heathlands coincided with the different grazing requirements of these habitats. Especially in large areas of conservation interest, we therefore encourage attempts to modify current wildlife management strategies to allow red deer to forage in open landscapes, which can contribute to the conservation of seminatural open habitats and also help to reduce damage in forest stands.

Abstract1. In agriculture, both valuable ecosystem services and unwanted ecosystem disservices can be produced by the same organism group. For example, small rodents can provide biological control through weed seed consumption but may also act as pests, causing crop damage.2. We studied the hypothesized causal relationships between ecosystem services (removal of weed seeds) and disservices (removal of wheat grains and crop damage) derived by small rodents (voles and mice) at multiple spatial scales. At the landscape scale, we studied the effects of landscape compositional and configurational heterogeneity on the abundance of voles and mice and their related ecosystem services and disservices along the former inner German border in east and west Germany. At the local scale, we studied how abundance and ecosystem functions are affected by management intensity (organic vs. conventional winter wheat), associated differences in crop characteristics and edge effects.3. Linear mixed-effects models and path analysis show that voles drove ecosystem disservices, but not ecosystem services, in agricultural fields. Daily wheat seed removal by voles was influenced by increasing wheat height and was almost three times higher than weed seed removal, which was not related to local-or landscape-scale effects.4. Abundance of voles and associated crop damage decreased with lower crop density and higher wheat height, which were associated with organic farming. Abundance of voles and crop damage were highest in conventional fields in west Germany. Synthesis and applications.As the ecosystem disservice of wheat seed consumption by small rodents must be considered mainly during crop sowing, management before crop harvest should focus on decreasing the pest potential of voles but not mice. Our results suggest that densities of voles and their ecosystem disservices could be reduced by having fields with low crop density and high wheat height, practices associated with organic farming. Surrounding landscapes with lowThis is an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium, provided the original work is properly cited.

Figure S1. Location of the sampling sites in heathlands (10) and grasslands (nine) at Grafenwöhr military training area (GTA) in Bavaria, Germany. Each site comprised four plots in heathlands and six plots in grasslands, respectively. Shaded areas indicate the occurrence of the Natura 2000 habitat types 4030, European dry heaths, and 6510, lowland hay meadows according to the draft of the Natura 2000 management plan (2013/2014) for the Site of Community Importance and Special Area of Conservation US-Truppenübungsplatz Grafenwöhr (DE6336301).

This is an open access article under the terms of the Creative Commons Attribution-NonCommercial License, which permits use, distribution and reproduction in any medium, provided the original work is properly cited and is not used for commercial purposes.

18Pollinators are sensitive to agricultural intensification at both local and landscape scales. High 19 configurational landscape heterogeneity due to small fields and a high amount of field 20 boundaries is hypothesized to enhance farmland biodiversity. Here, we investigated the 21 effects of organic vs. conventional farming in large-vs. small-scale agricultural landscapes on 22 wild bee communities and their floral resources to improve conservation schemes for 23 pollinators. We sampled bees in Central Germany at the field boundaries of 18 pairs of 24 conventionally and organically managed winter wheat fields along the former iron curtain 25 using pan traps and trap nests. Around traps, we surveyed species richness and flower cover 26 of insect-pollinated forbs. Compared to conventional farming, organic farming was related to 27 higher insect-pollinated forb species richness and flower cover in the field boundaries, 28 presumably due to the lack of herbicide use. Interestingly, small-scale agriculture did not 29 counteract the loss of floral resources under conventional management, as the difference to 30 organic management was even larger in Western small-scale agriculture. Organic farming, but 31 not small-scale agriculture, enhanced species richness of solitary bees, which is in line with 32 their small home ranges. In contrast, bumblebees benefitted only from small-scale agriculture, 33 which matches with their high dispersal ability. Despite similar levels of abundance and 34 diversity of trap-nesting bees in the two landscape types, brood cell parasitism was also 35 higher in small-scale agriculture. Both organic farming and small-scale agriculture directly 36 and indirectly supported different groups of wild bees, suggesting long-term benefits for 37 conservation. Agri-environment schemes should acknowledge the so far neglected benefits of 38 small-scale agriculture for biodiversity and its potential services. 39 40

Semi‐natural grasslands represent ecosystems with high biodiversity. Their conservation depends on the removal of biomass, for example, through grazing by livestock or wildlife. For this, spatially explicit information about grassland forage quantity and quality is a prerequisite for efficient management. The recent advancements of the Sentinel satellite mission offer new possibilities to support the conservation of semi‐natural grasslands. In this study, the combined use of radar (Sentinel‐1) and multispectral (Sentinel‐2) data to predict forage quantity and quality indicators of semi‐natural grassland in Germany was investigated. Field data for organic acid detergent fibre concentration (oADF), crude protein concentration (CP), compressed sward height (CSH) and standing biomass dry weight (DM) collected between 2015 and 2017 were related to remote sensing data using the random forest regression algorithm. In total, 102 optical‐ and radar‐based predictor variables were used to derive an optimized dataset, maximizing the predictive power of the respective model. High R2 values were obtained for the grassland quality indicators oADF (R2 = 0.79, RMSE = 2.29%) and CP (R2 = 0.72, RMSE = 1.70%) using 15 and 8 predictor variables respectively. Lower R2 values were achieved for the quantity indicators CSH (R2 = 0.60, RMSE = 2.77 cm) and DM (R2 = 0.45, RMSE = 90.84 g/m²). A permutation‐based variable importance measure indicated a strong contribution of simple ratio‐based optical indices to the model performance. In particular, the ratios between the narrow near‐infrared and red‐edge region were among the most important variables. The model performance for oADF, CP and CSH was only marginally increased by adding Sentinel‐1 data. For DM, no positive effect on the model performance was observed by combining Sentinel‐1 and Sentinel‐2 data. Thus, optical Sentinel‐2 data might be sufficient to accurately predict forage quality, and to some extent also quantity indicators of semi‐natural grassland.

Eutrophication through atmospheric nutrient deposition is threatening the biodiversity of semi-natural habitats characterized by low nutrient availability. Accordingly, local management measures aiming at open habitat conservation need to maintain habitat-specific nutrient conditions despite atmospheric inputs. Grazing by wild herbivores, such as red deer (Cervus elaphus), has been proposed as an alternative to mechanical or livestock-based measures for preserving open habitats. The role of red deer for nutrient dynamics in protected open habitat types, however, is yet unclear. Therefore, we collected data on vegetation productivity, forage removal, quantity of red deer dung and nutrient concentrations in vegetation and dung from permanent plots in heathlands and grasslands (eight plots à 225 m2 per habitat type) on a military training area inhabited by a large population of free-ranging red deer over one year. The annual nutrient export of nitrogen (N) and phosphorus (P) by red deer grazing was higher than the nutrient import through red deer excreta, resulting in an average net nutrient removal of 14 and 30 kg N ha−1 a−1 and 1.1 and 3.3 kg P ha−1 a−1 in heathlands and grasslands, respectively. Even when considering approximate local atmospheric deposition values, net nutrient depletion due to red deer grazing seemed very likely, notably in grasslands. Demonstrating that grazing by wild red deer can mitigate the effects of atmospheric nutrient deposition in semi-natural open habitats similarly to extensive livestock grazing, our results support the idea that red deer are suitable grazing animals for open habitat conservation.