Deposition and soil leaching in stands of Norway spruce and European Beech: Results from the Höglwald research in comparison with other European case studies

Stands of Norway spruce (Picea abies K.) and European beech (Fagus sylvatica L.) were investigated at the Höglwald research area, Southern Germany from 1985–1988 and from 1994–1997 in order to determine the effects of tree species on deposition and soil solution fluxes. The results were compared to 15 European case studies representing different deposition levels and site conditions. At the Höglwald site, which is characterised by a high nitrogen and a moderate sulphur load, throughfall deposition of nitrogen and sulphur compounds was about two-fold higher in spruce stands compared to beech stands. The differences in elemental input were clearly reflected in soil solution chemistry with a higher leaching of nitrate and sulphate in the spruce stands. The turnover of sulphur and nitrogen compounds induced a strong soil internal production of protons especially in the spruce stands. These results are in accordance with the other European case studies. Throughfall deposition and soil leaching of nitrogen and sulphur compounds was generally higher for spruce stands compared to beech stands. The species-related differences were mainly caused by dry deposition and were relatively small in remote areas. The consequences for the forest ecosystem itself and for the hydrosphere are discussed.     

The Höglwald field experiments – aims,concept and basic data

An outline is given of the aims and the concept of the interdisciplinary research work exerted at the Höglwald, an experimental field station in Southern Germany. The Höglwald lies in the temperate suboceanic broadleaf zone. Under natural conditions beech would prevail. The soils are mostly developed as Typic Hapludalfs, weakly aquic in the argillic horizon and strongly acidified in the topsoil. The investigations performed in stands of Norway spruce (Picea abies L. Karst) and European beech (Fagus sylvatica L.) were initiated in 1982/83. The main aims are (1) to test experimentally the soil acidification hypothesis predicting root damage by aluminium toxicity, (2) to study the effects of compensatory liming with and without artificially increased acid input, (3) to investigate comprehensively nitrogen cycling and turnover including studies about (a) N-gas fluxes, (b) root reactions to excess N-supply, (c) N-assimilation in the canopy, (d) effects of experimentally increased N-input on trees, soil and seepage water chemistry, and (e) N-turnover under natural and experimentally produced drought periods. Part of these investigations were also performed in a mixed stand of spruce and beech to show possible interactions of both species.     

Callose in roots of Norway spruce (Picea abies (L.) Karst.) is a sensitive parameter for aluminium supply at a forest site (Höglwald)

Under controlled environmental conditions in nutrient solution experiments induction of non-constitutive callose in roots has been shown to be a symptom of aluminium (Al) toxicity. In the present study roots of Norway spruce were sampled from a forest site where soil conditions had been modified by acidic irrigation and liming (Höglwald Experiment in Bavaria, Germany). A significant positive relationship was found between the callose content in short roots and the Al concentration in the soil solution, particularly if free Al, rather than total concentrations of soluble Al, were used for prediction. At the same sites root growth of Norway spruce was not affected by free Al concentrations in the range of 2.5 to 199 µM Al. The results show that also under field conditions a positive relationship between Al supply and callose content can be established. In Norway spruce callose content in roots is a much more sensitive parameter for Al supply than root growth.     

Evidence for a nutritional disorder of Oxalis acetosella L. on acid forest soils; II. Diagnostic field experiments and nutrient solution studies

Different field experiments were performed to discover the main factor(s) responsible for the poor leaf growth, moderate propagation and leaf chlorosis of Oxalis acetosella in an old Norway spruce stand with acid soil (Höglwald Forest). A previous study had suggested, that Ca (or Mg) deficiency or Mn toxicity could be involved.In a Main Diagnostic Field Experiment with an intact population, Ca and Mg were either applied as sulphate or carbonate to distinguish between nutritional and pH effects. Mn and Si were also applied to aggravate or overcome possible effects of Mn toxicity. Enhancement experiments with different amounts of CaSO₄ were conducted to investigate the Ca dose-effect relationship under field conditions. Additional trials with SrCO₃, BaCO₃ and NaHCO₃ had the goal to raise the soil pH without supply of nutrients.Greenhouse experiments with Oxalis acetosella supplemented the field studies by investigating the Ca and Mn dose-effect relationships under controlled conditions. Growth, vitality and nutrition of Oxalis were studied in a nutrient solution culture at pH 4.0 over a range of concentrations of Ca (20 to 5000 mol L_-1) and Mn (5 to 1000 mol L_-1) respectively. Furthermore, the effects of two contrasting ammonium/nitrate ratios were tested. The nutritional composition of the basal nutrient solution and the microclimate in the greenhouse were as far as possible adjusted to the environmental conditions of the plant in the Höglwald Forest.All these studies led to the conclusion, that the moderate growth and vitality of Oxalis in the Höglwald Forest was mainly due to an insufficient Ca supply, rather than an effect of Mg deficiency, low soil pH or Mn toxicity. The application of CaSO₄ caused a similar stimulation of the growth as CaCO₃. A clearly positive, close CaSO₄ dose-effect relationship was detected in field experiments as well as in the nutrient solution study. The same type of leaf chloroses as in the field was reproduced through low Ca nutrient solutions. Predominant ammonium nutrition may significantly impair Ca uptake.Oxalis acetosella displayed a relatively high leaf tissue tolerance of excessive Mn. There was no indication for a Mn-induced Ca deficiency in the Höglwald Forest. Enhanced Si uptake led to a partial vitalization of Oxalis; the reason for that remained unclear.     

Effects of plant diversity,N fertilization,and elevated carbon dioxide on grassland soil N cycling in a long‐term experiment

The effects of global environmental changes on soil nitrogen (N) pools and fluxes have consequences for ecosystem functions such as plant productivity and N retention. In a 13‐year grassland experiment, we evaluated how elevated atmospheric carbon dioxide (CO₂), N fertilization, and plant species richness alter soil N cycling. We focused on soil inorganic N pools, including ammonium and nitrate, and two N fluxes, net N mineralization and net nitrification. In contrast with existing hypotheses, such as progressive N limitation, and with observations from other, often shorter, studies, elevated CO₂ had relatively static and small, or insignificant, effects on soil inorganic N pools and fluxes. Nitrogen fertilization had inconsistent effects on soil N transformations, but increased soil nitrate and ammonium concentrations. Plant species richness had increasingly positive effects on soil N transformations over time, likely because in diverse subplots the concentrations of N in roots increased over time. Species richness also had increasingly positive effects on concentrations of ammonium in soil, perhaps because more carbon accumulated in soils of diverse subplots, providing exchange sites for ammonium. By contrast, subplots planted with 16 species had lower soil nitrate concentrations than less diverse subplots, especially when fertilized, probably due to greater N uptake capacity of subplots with 16 species. Monocultures of different plant functional types had distinct effects on N transformations and nitrate concentrations, such that not all monocultures differed from diverse subplots in the same manner. The first few years of data would not have adequately forecast the effects of N fertilization and diversity on soil N cycling in later years; therefore, the dearth of long‐term manipulations of plant species richness and N inputs is a hindrance to forecasting the state of the soil N cycle and ecosystem functions in extant plant communities.     

Forest herbs show species-specific responses to variation in light regime on sites with contrasting soil acidity: An experiment mimicking forest conversion scenarios

Forest conversion from native deciduous forests to coniferous stands has been performed in many European regions and resulted in dramatic shifts in understorey plant community composition. However, the drivers for changes in specific understorey plant species remained unclear.Here, we experimentally determine the species-specific effects of light availability and chemical soil characteristics, on the vegetative and regenerative performance of five herbaceous forest understorey plants. Topsoil samples from both spruce and deciduous stands at four locations, with two levels of soil acidity, were collected and used in a common garden experiment. Additionally, three different light levels were applied, i.e., ‘light deciduous’, ‘dark deciduous’ (extra light reduction during summer) and ‘evergreen’ (light reduction during winter). In a second experiment we evaluated the germination of two of these species against the acidity and tree species at the site of origin of the soil samples.The light regime affected both the vegetative and regenerative performance of the understorey species: compared to light deciduous, Anemone nemorosa had a significantly lower performance under the evergreen light regime, Convallaria majalis under dark deciduous and Luzula luzuloides and Galium odoratum under both light regimes. The vegetative performance was lower in soil from acid sites for the acid-sensitive species G. odoratum and Primula elatior. Differences between the soils sampled under deciduous or spruce stands had no effect on the vegetative, or the regenerative performance of these species. By contrast, the germination of L. luzuloides and P. elatior was higher in soils sampled in deciduous stands and in neutral sites.Species-specific responses in vegetative and regenerative performance of adult plants to a changed light regime and soil acidification could be a reason for the changed vegetation composition in converted stands. Also lower germination and establishment of forest understorey species in spruce stands could influence the species distribution after conversion.     

Deposition of ammonia to the forest floor under spruce and beech at the Höglwald site

Laboratory and field measurements of the flux of ammonia to forest floor canopies of spruce and beech stands at the Höglwald site in southern Bavaria are reported. Measurements were performed with an open chamber method. A linearity between ammonia concentration and ammonia flux from the atmosphere to the ground floor canopy was detected. Deposition of ammonia showed no saturation even at air concentrations up to 50 g NH₃ m^–3 air. Temperature, water content and the moss layer of the ground floor canopy had a minor influence on the deposition velocity in laboratory experiments. Deposition velocity of ammonia was higher to the spruce (1.3 cm s^–1), and limed spruce ground floor canopy (1.17 cm s^–1) compared to the beech stand (0.79 cm s^–1). In field studies, a diurnal course of the deposition velocity was detected with highest velocities in midday and minor during night times, but not in the climatic chamber. The flux of ammonia to the ground floor canopy was estimated of app. 10 kg N ha^–1 yr^–1 for the soil under spruce, 9 kg N ha^–1 yr^–1 for the limed spruce and 6 kg N ha^–1yr^–1 for the soil under beech. The fluxes are interpreted as fluxes from the atmosphere to the ground canopies of the stands.     

Effects of changes in tree species composition on water flow dynamics – Model applications and their limitations

Water-plant relations play a key role in the water cycling in terrestrial ecosystems. Consequently, changes in tree species composition may have distinct effects on the water retention capacity as well as on the pattern of streamflow generation. Such changes may result from modified interception properties and transpiration related to differences in canopy properties and root distribution. In order to evaluate the potential hydrological effects of the current silvicultural conversion from monocultural conifer stands into mixed or pure deciduous stands the hydrological model BROOK90 was applied to two forested upland catchments in Germany. The Rotherdbach catchment (9.4 ha, 93 yr-old Norway spruce) is situated in the Eastern Ore Mountains. The Schluchsee catchment (11 ha, 55-yr-old Norway spruce) is located in the higher altitudes of the Black Forest. The calibrated model is capable to describe rather well the temporal variation of streamflow but also the portions of the individual flow components. Data for a beech scenario were adapted for each site using a standard parameter set for deciduous trees provided by BROOK90. The annual discharge in the fictional beech stand at Rotherdbach is 30 to 50% higher compared to spruce with an increase of soil moisture and especially the slow streamflow components. This mainly results from low interception rates during winter time. In contrast, the spruce stand has a permanently higher interception rate. Effects of tree species conversion are moderate at Schluchsee. The annual discharge of a fictional beech stand at Schluchsee is 7 to 14% higher compared to spruce. There in contrast to Rotherdbach, effects of tree species conversion on soil moisture dynamics are small since vertical percolation in the highly permeable soil dominates and precipitation is abundant. Practical forestry will favorably establish mixed beech–spruce rather than pure beech stands. However, it is critical to simulate mixed stands with BROOK90. Therefore, a simple summation of model results from spruce and beech according to their respective area in a fictional mixed stand can only be a first approximation. Advanced hydrological simulation of mixed stand conditions should regard interactions of tree species and spatial parameter distribution. However, this is not yet feasible due to a distinct lack of information. As a consequence, there is a strong need to collect relevant hydrological and ecophysiological data in mixed stands in the future.     

Regeneration of mature Norway spruce stands: early effects of selective cutting and clear cutting on seepage water quality and soil fertility

The cutting of trees influences element turnover in the forest ecosystem. The reduction of plant uptake, as well as an increased mineralization and nitrification due to higher soil temperature and soil moisture, can lead to considerable losses of nutrients from the main rooting zone. This may result in a reduced soil fertility and a decrease in drinking water quality due to high nitrate concentrations in the seepage water. In Bavaria (Germany) selective cutting is preferred to clear cutting when initiating the regeneration of Norway spruce stands with European beech. This paper summarizes the early effects of both forest management practices on soil fertility and seepage water quality for three different sites. Shown are the concentrations of nitrogen and base cations in the seepage water as well as the water and ion fluxes during the first year after tree cut. Nutrient inputs decreased on thinned plots and even more at clear-cuts. Nitrate concentrations in the seepage water are hardly affected by moderate thinning; however, on clear-cuts, the nitrate concentration increases significantly, and base cations are lost from the upper mineral soil. This effect is less obvious at sites where a dense ground vegetation, which is able to take up excess nitrogen, exists.     

Effects of forest liming on soil processes

On the basis of a field experiment in Norway spruce with acid irrigation and compensatory liming of the soil surface (Höglwald, S-Bavaria), liming effects are described as lime dissolution rate, transformation of carbonate buffer to exchange buffer, time required for deacidification of soil and drainage water, mobilization of Cu and Pb, changes in soil organisms, humus decomposition, and nitrogen turnover. It was shown that lime dissolution followed an exponentially decreasing curve. 4 t ha^-1 dolomitic lime were dissolved within 6 years. Additional acid irrigation of 4 kmol H⁺ ha^-1 yr^-1 as sulphuric acid speeded up the lime dissolution to about 4 years. After dissolution of lime about 70% of Ca and about 30% of Mg, both originating from lime dissolution, are retained in the surface humus layer, loading the exchange buffer capacity there. Liming acted as a protection against acid irrigation but the extension of soil deacidification downwards proceeded slowly due to the high base neutralizing capacity of protonated functional groups of the organic matter. The main depth effect is caused by Mg translocation. A significant increase of organic Cu complexes occurred due to mobilization of water soluble humus decomposition products. The effect of liming on litter decomposing organisms is demonstrated with microorganisms, collembolae and earthworms regarding the abundance and the structure of dominance. It was shown that liming may induce unusually large changes in biocenoses of forest soils. The decay of surface humus accounted for 7.2 t ha^-1 or 23% of the store within 7 years. Within the same time span, liming caused a loss of about 170 kg N ha^-1 or 14% of the store of the surface humus layer. The nitrate concentration in the drainage water thus increased by about 50 to 60 mg NO₃ ^- L^-1. Site-specific conditions are discussed, which produce such negative liming effects as increased nitrate concentration of seepage, humus decay and heavy metal mobilization. Redistribution of tree roots, induction of boron deficiency and root rot are also considered. It is indicated that liming may aggravate the increasing problem of nitrate contamination of forest ground water resources which is associated with deposition of atmogenous nitrogen compounds. Some recommendations are given regarding forest practice.     

Soil seed banks of pure spruce (Picea abies) and adjacent mixed species stands

The soil seedbank of long living seeds of herbs, graminoids and shrubs can survive several decades in the soil and germinate after disturbances like windthrow or clearcutting. The main goal of this study was to evaluate the risk of weeds, which may limit the success of conversions of secondary pure spruce stands (Picea abies) to mixed species stands. In a first step, germination experiments were performed in the greenhouse on soil samples collected under adjacent pure spruce and mixed species stands (mainly mixtures of spruce and beech –Fagus sylvatica) on two different soil substrates (Flysch: nutrient rich, basic soil; Molasse: nutrient poor, acidic soil). Seedling density and species richness were higher on the nutrient rich soil on Flysch. Comparisons between seedlings that emerged from soil samples collected at the end of the vegetation period and in spring justify the statement of the hypothesis that mixed spruce-beech stands advance the transient seed bank while pure spruce stands stimulate the persistent seed bank. In a second step, the seed banks of different soil horizons down to 35 cm soil depth were studied in a multivariate statistical design for the most dominant species J. effusus, C. pallescens and R. idaeus, which are known to form long-term persistent seeds. Effects of bedrock material (Flysch, Molasse), species composition (pure spruce, mixed species) and treatment (control, nitrate) were tested. The total sum of these three species was significantly higher on Flysch than on Molasse. However, species composition indicated no significant differences, although there was a trend of higher amounts of germinating seeds under pure spruce. Nitrate treatments did not promote germination of viable buried seeds, indicating that the number of emerged seedlings is a realistic indicator of the seed bank density for the studied stands. It is concluded that overstorey tree species composition is not an important controlling factor for seed germination of the studied species after disturbances. The majority of emergents are the graminoids J. effusus and C. pallescens which were not present at all in the aboveground vegetation. Viable seeds were found down to 35 cm soil depth, although most seeds were concentrated in the upper 10 cm soil. Hence, care should be taken if management strategies create conditions that are generally favorable to germination. The success of forest regeneration or a conversion of pure spruce to mixed species stands could be endangered by any disturbance, which causes an immediate increase of light levels.     

The soil fauna community in pure and mixed stands of beech and spruce of different age: trophic structure and structuring forces

This study investigates the response of the soil fauna community to replacement of beech by spruce or by mixed stands of beech and spruce. Stands of different age were investigated in a factorial design with the factors tree species (beech and spruce) and stand age (30 and 120 yr). The input of leaf/needle litter did not differ significantly between the study sites. By contrast, the amount of organic matter in upper soil layers (L/F, H/Ah) of spruce forests strongly exceeded that of beech forests particularly in mature stands. The increase in organic matter in spruce stands was not associated by an increase in the amount of microbial biomass. Biomass of eight (bacterivorous, fungivorous and omnivorous nematodes, enchytraeids, earthworms, isopodes, mycetophilid and cecidomyiid Diptera) of the twelve microbi‐detritivorous soil animal groups studied was significantly increased in beech forests; only that of one group (elaterid beetles) was increased in spruce forests and three groups did not respond significantly (collembolans, oribatid mites, sciarid Diptera). This indicates that in the forests studied neither habitat space (amount of organic matter in L/F and H/Ah layers) nor the amount of microbial biomass controlled microbi‐detritivores. Rather, the quality of litter materials and the concentration of microbial biomass therein appeared to be most important. Herbivores and predators also were favoured by beech: the biomass of one (rhizophagous nematodes) of the three herbivorous groups studied were significantly increased in beech stands and none in spruce stands; the biomass of four (predatory nematodes, centipedes, carabid and cantharid beetles) of the seven carnivorous groups studied were increased in beech stands, none in spruce stands. Generally, the biomass ratio between prey and predators was at a minimum in mature beech and mixed stands indicating more intense top‐down control in these forests. Overall, the study documents that replacement of beech by spruce strongly alters the soil food web. Mixed stands were more similar to spruce stands in respect to the biomass of soil animal groups but predator–prey interactions appeared to be more similar in mature beech and mixed stands. Differences between tree species usually were more pronounced in 120 compared to 30 yr old stands indicating that the development of stand characteristics is slow.     

Ion concentrations and fluxes of seepage water before and after clear cutting of Norway spruce stands at Ballyhooly, Ireland, and Höglwald, Germany

Ion concentrations and fluxes in seepage water (below the main rooting zone) were compared before and after clear cutting at two similar long-term experimental Norway spruce forest plots. While Ballyhooly (Ireland) was influenced by sea salt deposition, Höglwald (Germany) received high nitrogen (N) deposition. These differences were reflected in seepage water concentrations with sodium (Na⁺) and chloride (Cl^–) dominating at Ballyhooly and high nitrate (NO₃ ^?) and aluminium concentrations at Höglwald. Following clear cutting of the forest plots, NO₃ ^? concentrations peaked (Ballyhooly: 2018 μmol_c L^?1, Höglwald: 2595 μmol_c L^?1). Moreover, at Ballyhooly, NO₃ ^? concentrations and fluxes were continuously elevated for ~1.5 years. At Höglwald, the clear cut plot, which was replanted with spruce and beech saplings, exhibited periodically elevated NO₃ ^? concentrations with two distinct peaks. However, low concentrations, compared to the control (uncut) plot, were also observed. Further, at Höglwald a plot with a pre-existing dense natural regeneration of Norway spruce exhibited much lower NO₃ ^? concentrations before and after clear cutting. Nonetheless, NO₃ ^? concentrations following clear cut at both sites were elevated at least periodically above European drinking water standards (50 mg L^?1). An important prerequisite for NO₃ ^? leaching is that forests are N saturated or at least not N-limited; consequently chronic elevated N deposition may lead to increased deterioration of seepage water quality across Europe following forest disturbances (harvesting, windthrow, insect attacks). Clear cutting at Ballyhooly was responsible for significant element loss, especially of potassium, N and calcium, while magnesium loss was compensated by high sea salt inputs. At Höglwald the contamination of seepage water with NO₃ ^? has been the main problem for more than 20 years at the mature stand. A pre-existing regeneration can help to reduce NO₃ ^? and cation leaching after cutting.     

Greater accumulation of litter in spruce (Picea abies) compared to beech (Fagus sylvatica) stands is not a consequence of the inherent recalcitrance of needles

Background and aims

Replacement of beech by spruce is associated with changes in soil acidity, soil structure and humus form, which are commonly ascribed to the recalcitrance of spruce needles. It is of practical relevance to know how much beech must be admixed to pure spruce stands in order to increase litter decomposition and associated nutrient cycling. We addressed the impact of tree species mixture within forest stands and within litter on mass loss and nutritional release from litter.

Methods

Litter decomposition was measured in three adjacent stands of pure spruce (Picea abies), mixed beech-spruce and pure beech (Fagus sylvatica) on three nutrient-rich sites and three nutrient-poor sites over a three-year period using the litterbag method (single species and mixed species bags).

Results

Mass loss of beech litter was not higher than mass loss of spruce litter. Mass loss and nutrient release were not affected by litter mixing. Litter decay indicated non-additive patterns, since similar remaining masses under pure beech (47%) and mixed beech-spruce (48%) were significantly lower than under pure spruce stands (67%). Release of the main components of the organic substance (C_org, N_tot, P, S, lignin) and associated K were related to mass loss, while release of other nutrients was not related to mass loss.

Conclusions

In contradiction to the widely held assumption of slow decomposition of spruce needles, we conclude that accumulation of litter in spruce stands is not caused by recalcitrance of spruce needles to decay; rather adverse environmental conditions in spruce stands retard decomposition. Mixed beech-spruce stands appear to be as effective as pure beech stands in counteracting these adverse conditions.     

Neighborhood effects on soil properties,mycorrhizal attributes,tree growth,and nutrient status in afforested zones

A harmonious interspecies relationship is the key to the success of mixed afforestation. This study was conducted to assess the responses of afforestation species to their neighboring trees. We examined five types of stands—monocultures of Chinese pine (Pinus tabuliformis), black locust (Robinia pseudoacacia), sea‐buckthorn (Hippophae rhamnoides), and two mixtures (Chinese pine × black locust mixture and Chinese pine × sea‐buckthorn mixture)—in the Loess Plateau, northwestern China. The height and diameter at breast height of each tree species were measured, and rhizosphere soil, shoot, and root were sampled. In monocultures, black locust was taller than Chinese pine and sea‐buckthorn, while the height of Chinese pine and sea‐buckthorn was similar. In mixtures, Chinese pine grew better with sea‐buckthorn than alone as a result of modified soil properties and plant nutrition, but not with black locust. When Chinese pine was used as neighbors, it affected the level of arbuscular mycorrhizal (AM) colonization of black locust, soil properties and AM fungal spore density of black locust and sea‐buckthorn, but did not significantly affect their growth. Our results suggest that the reciprocal effects between tree species in mixture are not symmetric, and thus planning for efficient mixed afforestation requires knowledge of species‐specific growth rate, nutrient requirements, and species interactions.     

Changes in soil properties after afforestation of former intensively managed soils with oak and Norway spruce

In many European countries, surplus agricultural production and ecological problems due to intensive soil cultivation have increased the interest in afforestation of arable soils. Many environmental consequences which might rise from this alternative land-use are only known from forest establishment on less intensively managed or marginal soils. The present study deals with changes in soil properties following afforestation of nutrient-rich arable soils. A chronosequence study was carried out comprising seven Norway spruce (Picea abies (Karst.) L.) and seven oak (Quercus robur L.) stands established from 1969 to 1997 on former horticultural and agricultural soils in the vicinity of Copenhagen, Denmark. For comparison, a permanent pasture and a ca. 200-year-old mixed deciduous forest were included. This paper reports on changes in pH values, base saturation (BS_eff), exchangeable calcium, soil N pools (N_min contents), and C/N ratios in the Ap-horizon (0–25 cm) and the accumulated forest floor. The results suggest that afforestation slowly modifies soil properties of former arable soils. Land-use history seems to influence soil properties more than the selected tree species. An effect of tree species was only found in the forest floor parameters. Soil acidification was the most apparent change along the chronosequence in terms of a pH decrease from 6 to 4 in the upper 5 cm soil. Forest floor pH varied only slightly around 5. Nitrogen storage in the Ap-horizon remained almost constant at 5.5 Mg N ha^–1. This was less than in the mineral soil of the ca. 200-year-old forest. In the permanent pasture, N storage was somewhat higher in 0–15 cm depth than in afforested stands of comparable age. Nitrogen storage in the forest floor of the 0–30-year-old stands increased in connection with the build-up of forest floor mass. The increase was approximately five times greater under spruce than oak. Mineral soil C/N ratios ranged from 10 to 15 in all stands and tended to increase in older stands only in 0–5 cm depth. Forest floor C/N ratios were higher in spruce stands (26.4) as compared to oak stands (22.7). All stands except the youngest within a single tree species had comparable C/N ratios.     

Four years continuous record of CH4-exchange between the atmosphere and untreated and limed soil of a N-saturated spruce and beech forest ecosystem in Germany

In order to gain information about seasonal and interannual variations of CH₄-fluxes at a spruce control site, a limed spruce site and a beech site of the Höglwald Forest, Bavaria, Germany, complete annual cycles of CH₄-exchange between the soil and the atmosphere with 2-hourly resolution were followed for 4 consecutive years. The ranges of CH₄ fluxes observed for the different sites were: +12.4 to –69.4 g CH₄ m^–2 h^–1 (spruce control site), +11.7 to –51.4 g CH₄ m^–2 h^–1 (limed spruce site), and –4.4 to –167.3 g CH₄ m^–2 h^–1 (beech site). Lowest rates of atmospheric CH₄-uptake or even a weak net-emission of CH₄ by the soils were observed during winter/spring times, whereas highest rates of CH₄-uptake were always found in summer/spring. Over the entire observation period of 4 years, mean CH₄-uptake rates were –1.82 kg CH₄-C ha^–1 yr^–1 at the spruce control site, –1.31 kg CH₄-C ha^–1 yr^–1 at the limed spruce site, and –4.84 kg CH₄-C ha^–1 yr^–1 at the beech site. The results obtained in this study demonstrate that in view of the huge interannual variations in CH₄-fluxes of approx. 1 kg CH₄-C ha^–1 yr^–1, multiple year measurements of CH₄-fluxes are necessary to accurately characterize the sink strength of temperate forest for atmospheric CH₄. By comparison of CH₄-fluxes measured at the spruce control site and the limed spruce site, a significant negative effect of forest floor liming on CH₄-uptake could be demonstrated. Compared to the spruce stand, the beech stand showed on average approx. 3 times higher rates of atmospheric CH₄-uptake, most likely due to pronounced differences between both sites with regard to the organic layer structure and bulk density of the mineral soil. Regression analysis between CH₄-fluxes and environmental parameters revealed that at all sites the dominating factors regulating temporal variations of CH₄ fluxes were soil moisture and soil temperature. Field measurements of CH₄ concentrations in the soil profile and laboratory measurements of CH₄-oxidation and CH₄-production activity on soil samples taken from different soil depths showed that the CH₄-flux at the Höglwald Forest sites is the net-result of simultaneous occurring production and consumption of CH₄ within the soil. Highest CH₄-oxidation activity was found in the uppermost centimeters of the mineral soil, whereas highest potential CH₄-production activity was found in the organic layer.     

Inorganic soil nitrogen under grassland plant communities of different species composition and diversity

We measured aboveground plant biomass and soil inorganic nitrogen pools in a biodiversity experiment in northern Sweden, with plant species richness ranging from 1 to 12 species. In general, biomass increased and nitrate pools decreased with increasing species richness. Transgressive overyielding of mixed plant communities compared to the most productive of the corresponding monocultures occurred in communities with and without legumes. N₂-fixing legumes had a fertilizing function, while non-legumes had a N retaining function. Plant communities with only legumes had a positive correlation between biomass and soil nitrate content, whereas in plant communities without legumes they were negatively correlated. Both nitrate and ammonium soil pools in mixed non-legume communities were approximately equal to the lowest observed in the corresponding monocultures. In mixed legume/non-legume communities, no correlation was found for soil nitrate with either biomass or legume biomass as percentage of total biomass. The idea of complementarity among species in nitrogen acquisition was supported in both pure non-legume and mixed non-legume/legume communities. In the latter, however, facilitation through increased nitrogen availability and retention, was probably dominating. Our results suggest that diversity effects on biomass and soil N pools through resource use complementarity depend on the functional traits of species, especially N₂ fixation or high productivity.     

Evidence for a nutritional disorder of Oxalis acetosella L. on acid forest soils; I. Control situation and effects of dolomitic liming and acid irrigation

This paper reports on the mean cover, biometric and vitality parameters and mineral nutrient status of wood-sorrel (Oxalis acetosella L.) on the control and experimental plots of a mature Norway spruce stand on acid soil (Höglwald Experiment).On the control plot Oxalis showed relatively poor leaf growth and chloroses on young and older leaf blades. Mineral analyses in the Höglwald and comparative studies on other forest sites indicated a very good to good supply of N and P, a moderate Ca nutrition and very high Mn concentrations of the leaves.The input of dolomitic lime led to a drastic propagation and vitalization of the wood-sorrel. The long-lasting effect of treatment was independent of pH dynamics in the root zone. The leaf chloroses of Oxalis disappeared completely after that treatment. The level of Ca and Mg supply clearly improved, whereas the concentrations of P, N, Mn and Zn diminished. Surprisingly, the K nutrition also improved after liming.Oxalis reacted to acid irrigation (sulfuric acid; pH 2.7-2.8) in comparison to normal watering (pH 5.0-5.5) with a continuous decrease in cover, but without visible leaf necroses. The concentrations of total sulphur, SO₄-S and S_org were raised, while the concentrations of Ca, Mn and Zn of the leaf blades decreased. After termination of acid input a recovery of nutrition occurred and a slow recolonization began. Preceding lime application prevented the growth-inhibiting effect of the acid irrigation.The study leads to the working hypothesis, that Ca nutrition may be the controlling factor for leaf growth and vegetative propagation of Oxalis in the Höglwald Experiment.     

Plant-soil feedback in spruce (Picea abies) and mixed spruce-beech (Fagus sylvatica) stands as indicated by dendrochemistry

Ten pairs of secondary pure spruce (Picea abies) and adjacent mixed spruce-beech (Fagus sylvatica) stands on comparable sites were selected on two different bedrocks for soil formation (Flysch: nutrient rich and high soil pH; Molasse: poor nutrient supply and acidic) to study how an admixture of beech to spruce stands affects nutrient cycling and consequently soil chemistry. Soil analyses indicated accumulation of Ca under the mixed stands while the top soil under pure spruce was acidified. It was hypothesized that changes of soil chemical properties due to species composition over the last six decades are reflected in the stem wood of spruce. Three healthy dominant spruce trees per plot were selected for coring. Cores were crossdated and half-decadal samples were analyzed for Ca, Mg, Mn and Al. Calcium and Mg concentrations in stem wood of spruce were significantly higher for the pure spruce than for the mixed stands in spite of lower Ca and Mg stores in the soil. We assume that acidification caused by pure spruce mobilized these cations temporarily, increasing soil solution contents and consequently stem wood concentrations. It was possible to reconstruct soil pH from the element ratios Ca/Al (pure stands) and Ca/Mg (mixed stands), since these ratios in the stem wood of the last half-decade did correlate with soil pH for selected soil depths. Reconstructed soil pH showed a decline over the last 60 years under both species compositions due to accumulation of base cations in the increasing biomass. Comparisons of reconstructed soil pH in 0–5 and 10–20 cm soil depth indicated more pronounced top soil acidification (lower soil pH in 0–5 cm) by spruce on the nutrient rich soil (Flysch) than on the acidic soil (Molasse). However, admixture of beech caused higher pH values in 0–5 cm than in 10–20 cm soil depth on Flysch due to the observed Ca-pump effect of beech (uptake of Ca from deeper soil horizons).