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In the present study we analyzed the combined effects of management (grazing, mowing, prescribed burning, sod-cutting) and atmospheric deposition on N and P budgets of heathland ecosystems (Lüneburger Heide nature reserve; N Germany). We hypothesize that management measures such as grazing and mowing can accelerate a deposition-induced imbalance of N and P pools as a result of a disproportionally high output of P. We analyzed management and deposition affected input-output flows of N and P and related them to changes in the nutritional status of Calluna vulgaris 5 years after treatment application. We found that grazing and mowing caused the highest net loss of P due to high P concentrations in the aboveground biomass. In contrast, prescribed burning only slightly affected P pools, as P remained in the system due to ash deposition. Management-mediated effects on N and P pools were mirrored in the nutritional status of Calluna vulgaris: at the grazed and mown sites, the P content of current season's shoots significantly decreased within 5 years after treatments, whereas the N content remained unchanged. We conclude that grazing and mowing can accelerate declining availability of P and, thus, accelerate a deposition-induced shift from N- to P-limited plant growth in the medium term. In the face of ongoing atmospheric N loads management schemes need to combine high- and low-intensity measures to maintain both a diverse structure and balanced nutrient budgets in the long term. © 2009 Springer Science+Business Media, LLC. In the present study we analyzed the combined effects of management (grazing, mowing, prescribed burning, sod-cutting) and atmospheric deposition on N and P budgets of heathland ecosystems (Lüneburger Heide nature reserve; N Germany). We hypothesize that management measures such as grazing and mowing can accelerate a deposition-induced imbalance of N and P pools as a result of a disproportionally high output of P. We analyzed management and deposition affected inputoutput flows of N and P and related them to changes in the nutritional status of Calluna vulgaris 5 years after treatment application. We found that grazing and mowing caused the highest net loss of P due to high P concentrations in the aboveground biomass. In contrast, prescribed burning only slightly affected P pools, as P remained in the system due to ash deposition. Management-mediated effects on N and P pools were mirrored in the nutritional status of Calluna vulgaris: at the grazed and mown sites, the P content of current season’s shoots significantly decreased within 5 years after treatments, whereas the N content remained unchanged. We conclude that grazing and mowing can accelerate declining availability of P and, thus, accelerate a deposition-induced shift from N- to P-limited plant growth in the medium term. In the face of ongoing atmospheric N loads management schemes need to combine high- and low-intensity measures to maintain both a diverse structure and balanced nutrient budgets in the long term.

Questions: What effect does sheep grazing have on the nutrient budgets of heathlands? Can grazing compensate for atmospheric nutrient loads in heathland ecosystems? What are the conclusions for heathland management? Location: Lüneburg Heath, NW Germany. Methods: During a one-year grazing experiment (stocking rate 1.1 sheep/ha) nutrient balances for N, Ca, K, Mg and P were calculated by quantifying input rates (atmospheric deposition, sheep excrement) and output rates (biomass removal, leaching). Results: Atmospheric nutrient deposition amounted to 22.8 kg.ha1.a1 for N and

Summary 1. Atmospheric nutrient deposition has contributed to widespread changes in heathlands throughout Europe. As a consequence, management is now being considered as a potential tool with which to compensate for increased nutrient loads. Currently, only limited information is available on the extent to which management measures could compensate for atmospheric nutrient deposition. We hypothesized that low-intensity management measures are not sufficient to counterbalance current nutrient inputs, particularly of nitrogen (N). 2. In order to improve heathland management schemes, we evaluated the effectiveness of different management measures in reducing the impact of ongoing atmospheric nutrient loads. We compared the effects of mowing, prescribed burning (low-intensity management) and sod-cutting (high-intensity management) on heathland nutrient budgets [N, calcium (Ca), potassium (K), magnesium (Mg) and phosphorus (P)] in the Lueneburg Heath nature reserve (north-west Germany). Nutrient balances were calculated by analysing the present-day input, the output as a result of the removal of biomass/humus horizons, and changes in leaching rates. 3. Nutrient losses by increased leaching following management measures were negligible compared with nutrient losses caused by the removal of above-ground biomass or humus horizons. The total quantities of nutrients removed by sod-cutting were equivalent to between 37 and 176 years of atmospheric input (for N, 89 years). 4. In contrast, the quantities of N removed by mowing and prescribed burning were equivalent to only 5 years of atmospheric input. Thus, heathlands subjected to such treatments will accumulate N in the long term. In addition, output‐input ratios for phosphorus (P) exceeded those for N in the mowing and sod-cutting experiments. It is therefore likely that heathlands currently (co-) limited by N will shift to being more Plimited in the long term. This will promote species that are well adapted to P-limited sites (e.g. Molinia caerulea ). 5. Synthesis and applications . This study shows that low-intensity management cannot compensate for atmospheric N loads in the long term. Consequently, high-intensity management measures are an indispensable tool in preserving a long-term balanced N budget in heathlands. In order to maintain a diverse structure, managers need to combine low- and high-intensity management measures. Prescribed burning proved to be the best means of avoiding an increasing P shortage, because this measure causes very low P outputs.

Question: Can prescribed winter burning compensate atmospheric nutrient loads for dry heathlands? What effects does prescribed burning have on nutrient balances, particularly as regards the limiting nutrients N and P? Location: Lueneburg Heath, NW Germany. Methods: In two burning experiments (in 10/15 year old Calluna-stands) nutrient balances (for N, Ca, K, Mg, P) were calculated by analysing nutrient inputs (atmospheric deposition, ash deposition), nutrient stores (above-ground biomass, organic horizon) and nutrient outputs (biomass combustion, leaching). Results: Atmospheric nutrient deposition amounted to 22.8 kg.ha -1.a -1 for N and < 0.5 kg.ha -1.a -1 for P. Nutrient stores in the above-ground biomass were 95/197 kg.ha -1 for N and 5/13 kg.ha -1 for P (first/second experiment, respectively). From these stores 90/53% (for N) and 25/14% (for P) were removed by burning. Effects of leaching on nutrient balances were low. In the first two years after burning, leaching rates of N increased by about 4/6 kg.ha -1, whereas leaching rates of P did not change significantly. Input/output-ratios showed that prescribed burning leads to positive nutrient balances for N, Ca and Mg in the long term. For example, the amounts of N removed by prescribed burning are equivalent to ca. five years of atmospheric inputs. Applied in ten-year cycles, this measure alone cannot prevent N accumulation in the long term. Conclusion: Regarding 10/15 year old Calluna-heaths, we assume that prescribed burning cannot compensate for atmospheric N inputs, thus making long-term changes in the nutritional state inevitable. Therefore, prescribed burning should be applied in combination with high-intensity management measures.

Atmospheric nutrient deposition has contributed to widespread changes in sensitive seminatural ecosystems throughout Europe. For an understanding of underlying processes it is important to quantify input-output flows in relation to ongoing atmospheric inputs and current management strategies. In this study we quantified losses of N, P, Ca, Mg, and K via leaching in heathland ecosystems (Lüneburger Heide, NW Germany) as a function of current deposition rates and different management measures (mowing, prescribed burning, choppering, sod-cutting) which aim to prevent shrub and tree encroachment. Leaching was only moderately related to atmospheric input rates, indicating that leaching was mostly affected by internal turnover processes. Leaching significantly increased for most of the nutrients after the application of management measures, particularly in the choppered and sod-cut plots. However, atmospheric nutrient inputs exceeded leaching outputs for most of the nutrients, even in the plots subjected to management. Despite high deposition rates (20-25 kg N ha -1year-1), retention of atmospheric N input ranged between 74% and 92% in the control plots. In the treated plots, N retention decreased to 59-80%. However, in the study area mean N leaching in the controls has almost doubled since 1980 and currently amounts to 3.7 kg ha-1year -1, indicating an early stage of N saturation. Our study provides evidence that leaching did not compensate for atmospheric nutrient deposition, particularly as regards N. Management, thus, will be an indispensable tool for the maintenance of the low-nutrient status as a prerequisite for the long-term preservation of heathland ecosystems. © 2007 Springer Science+Business Media B.V.

Heathlands are endangered by both atmospheric nutrient deposition and natural succession. High-intensity management measures are considered necessary as low-intensity measures (e.g. mowing, prescribed burning) are not able to compensate for atmospheric nutrient loads. Choppering (i.e. the near-complete removal of the O-layer) has several advantages over sod-cutting, including less waste matU:/ES/DTD501/Bioc/3186erial, faster vegetation recovery and lower costs. This raises the question addressed in this study as to the extent to which choppering and sod-cutting affect nutrient budgets in dry heath-lands. We compared the quantities of N, Ca, K, Mg, and P removed by choppering and sod-cutting in the Lueneburg Heath (NW Germany). Nutrient balances were calculated by analysing atmospheric inputs, elevated leaching rates following management, and output due to the removal of above-ground biomass and humus horizons. Nutrient loss was particularly high after removal of O- and A-horizons. in contrast, increased leaching after management was of minor importance for nutrient budgets. Although considerably more nutrients were removed by sod-cutting than by choppering (e.g. N: 1712/1008 kg ha(-1)), nutrient output by choppering was still sufficient to compensate for 60.7 years of net N-input. Choppering was able to remove more N per volume unit than sod-cutting due to higher N-contents in the organic layer than in the A-horizon. For this reason, choppering is more economical than sod-cutting and, thus, should be considered the preferable method at sites not dominated by Molinia caerulea. A combination of high-intensity measures with prescribed burning would appear to be suitable as this would ensure more selective removal of N. (c) 2006 Elsevier Ltd. All rights reserved. Heathlands are endangered by both atmospheric nutrient deposition and natural succession. High-intensity management measures are considered necessary as low-intensity measures (e.g. mowing, prescribed burning) are not able to compensate for atmospheric nutrient loads. Choppering (i.e. the near-complete removal of the O-layer) has several advantages over sod-cutting, including less waste matU:/ES/DTD501/Bioc/3186erial, faster vegetation recovery and lower costs. This raises the question addressed in this study as to the extent to which choppering and sod-cutting affect nutrient budgets in dry heath-lands. We compared the quantities of N, Ca, K, Mg, and P removed by choppering and sod-cutting in the Lueneburg Heath (NW Germany). Nutrient balances were calculated by analysing atmospheric inputs, elevated leaching rates following management, and output due to the removal of above-ground biomass and humus horizons. Nutrient loss was particularly high after removal of O- and A-horizons. in contrast, increased leaching after management was of minor importance for nutrient budgets. Although considerably more nutrients were removed by sod-cutting than by choppering (e.g. N: 1712/1008 kg ha(-1)), nutrient output by choppering was still sufficient to compensate for 60.7 years of net N-input. Choppering was able to remove more N per volume unit than sod-cutting due to higher N-contents in the organic layer than in the A-horizon. For this reason, choppering is more economical than sod-cutting and, thus, should be considered the preferable method at sites not dominated by Molinia caerulea. A combination of high-intensity measures with prescribed burning would appear to be suitable as this would ensure more selective removal of N. (c) 2006 Elsevier Ltd. All rights reserved.