High organic carbon stock in a karstic soil of the Middle-European Forest Province persists after centuries-long agroforestry management

Little is known on soil organic carbon (SOC) stocks in karst areas worldwide, although many of them have seen long-term application of agroforestry systems with a potential for carbon sequestration. Therefore, our study aimed to assess landscape-level SOC concentration and stock in the Silica Plateau, a part of the Slovak Karst Biosphere Reserve located in the Western Carpathians (Slovakia) with a centuries-long agroforestry record. The most represented local soil units are Chromi-Rendzic Leptosols and Chromic Cambisols with clayey loam texture, C/N ratio 9–12, and $ {\text{pH}}_{{{\text{H}}_{2} {\text{O}}}} $ 6.6–6.2 in their organo-mineral surface horizons. Mull surface humus form prevails under mixed forest stands dominated by hornbeam (Carpinus betulus L.), oak (Quercus petraea L.), and beech (Fagus sylvatica L.). A total of 2,700 soil samples were collected from 150 soil pits. Both SOC concentrations and stocks were determined for the 0–60?cm mineral soil layer. Soil stoniness was accounted for by means of electrical resistivity tomography. According to the analysis of covariance, cropland SOC concentration (0.026?g?g^?1) is significantly lower compared to forestland (0.040?g?g^?1) and pastureland (0.041?g?g^?1) (P?<?0.01). During the period of 130?years after forest clearing, cropland SOC stock has been reduced at an exponential decay rate of ca 0.002?year^?1, while the SOC stock in pastureland has increased following land use change from cropland by approximately 30% during the same period of time. Irrespective of land use history, overall SOC stock is high reaching on average 207.4?Mg?ha^?1, out of which 66% are stored within 0–30?cm and 34% within 30–60?cm soil layers.     

Phosphorus supply and cycling at long-term forest monitoring sites in Germany

In this study, the supply and input–output balances of phosphorus (P) were investigated for a 10-year-period at 85 long-term monitoring sites in German forest ecosystems under the European Level II programme. These sites encompass 23 European beech (Fagus sylvatica L.) stands, 9 oak stands comprised of common oak (Quercus robur L.) and/or sessile oak (Quercus petraea Liebl.), 20 Scots pine (Pinus sylvestris L.) and 33 Norway spruce (Picea abies H.Karst.) stands. We quantified P concentrations in needles and leaves, P inputs from the atmosphere, P outputs through leaching and harvesting, and total P in the soil and humus layers. The P concentrations in European beech leaves from two sites (>1 mg P g⁻¹ dry weight), and in Norway spruce needles from four sites (>1.2 mg P g⁻¹ dry weight), were deficient over several years. In contrast, the oak and Scots pine sites were well supplied with P. When P removal through harvesting was disregarded, P balances were positive or stable (median 0.21 kg P ha⁻¹ a⁻¹). With harvesting, balances were mostly negative (median −0.35 kg P ha⁻¹ a⁻¹), with long-term P removal from the forest ecosystems.     

Carbon and nitrogen dynamics in topsoils along forest conversion sequences in the Ore Mountains and the Saxonian lowland, Germany

Carbon and nitrogen stocks and their medium-term and readily decomposable fractions in topsoils were compared in relation to soil microbial biomass and activity along sequences from coniferous to deciduous stands. The study was carried out in the Ore Mountains and the Saxonian lowland, representing two typical natural regions in Saxony, Germany. In accordance with current forest conversion practices, the investigation sites represent different stands: mature conifer stands of Norway spruce (Picea abies (L.) Karst.) and Scots pine (Pinus sylvestris L.) (type A); Norway Scots spruce and pine with advanced plantings of European beech (Fagus sylvatica L.) or European beech/Common oak (Quercus petreae Liebl.) (type B); and mature deciduous stands of European beech and European beech/Common oak (type C). The investigated forest sites can be grouped into three silvicultural situations according to the development from coniferous stands to advanced plantings and finally mature deciduous forests (chronosequence A–B–C). The organic layer (L, F and H horizons) and uppermost mineral soil (0–10 cm) were analysed for potential C mineralisation, microbial biomass, concentrations of total C and N (TOC and TN) and for medium-term and readily decomposable C and N fractions, obtained by hot- and cold-water extraction respectively. The results showed an increase in organic layer thickness and mass as well as TOC and TN stocks along the forest sequences in the lowland. Yet, underplanted sites with two storeys revealed higher organic layer mass as well as TOC and TN stocks as compared to coniferous and deciduous stands. Stocks of hot- and cold-water-extractable C and N in relation to microbial biomass and its activity revealed a high turnover activity in deeper organic horizons of deciduous forests compared to coniferous stands. The stand-specific differentiation is discussed in relation to microbial biomass, litter quantity and quality and forest structure, but also with respect to the site-specific climatic factors and water budget as well as liming and fly-ash impacts. Results indicate higher dynamics in deciduous stands in the lowland especially during the initial turnover phase. The elevated microbial activity in deeper organic horizons of deciduous litter-influenced sites in spring is discussed as a specific indicator for long-term C sequestration potential as besides C mineralisation organic compounds are humified and thus, can be stored in the organic layer or in deeper soil horizons. Due to liming activities, stand-specific effects on organic matter turnover dynamics have evened out today in the Ore mountain region, but will presumably occur again once base saturation decreases. Here, the stand-specific effect on microbial biomass can currently be seen again as C_mic in the L horizon increased from spruce to beech. Our study sites in the lowland revealed no significant fly-ash impact. Differences between sites were evaluated by calculating the discriminance function. TOC and TN as well as medium-term degradable C and N were defined in this study as indicators for turnover dynamics along forest conversion sites.     

Stocks and dynamics of soil organic carbon and coarse woody debris in three managed and unmanaged temperate forests

The effect of forest conservation on the organic carbon (C) stock of temperate forest soils is hardly investigated. Coarse woody debris (CWD) represents an important C reservoir in unmanaged forests and potential source of C input to soils. Here, we compared aboveground CWD and soil C stocks at the stand level of three unmanaged and three adjacent managed forests in different geological and climatic regions of Bavaria, Germany. CWD accumulated over 40–100 years and yielded C stocks of 11 Mg C ha^?1 in the unmanaged spruce forest and 23 and 30 Mg C ha^?1 in the two unmanaged beech–oak forests. C stocks of the organic layer were smaller in the beech–oak forests (8 and 19 Mg C ha^?1) and greater in the spruce forest (36 Mg C ha^?1) than the C stock of CWD. Elevated aboveground CWD stocks did not coincide with greater C stocks in the organic layers and the mineral soils of the unmanaged forests. However, radiocarbon signatures of the O _e and O _a horizons differed among unmanaged and managed beech–oak forests. We attributed these differences to partly faster turnover of organic C, stimulated by greater CWD input in the unmanaged forest. Alternatively, the slower turnover of organic C in the managed forests resulted from lower litter quality following thinning or different tree species composition. Radiocarbon signatures of water-extractable dissolved organic carbon (DOC) from the top mineral soils point to CWD as potent DOC source. Our results suggest that 40–100 years of forest protection is too short to generate significant changes in C stocks and radiocarbon signatures of forest soils at the stand level.     

Growth and mortality in underplanted tree seedlings in response to variations in canopy closure of Norway spruce stands

Six broadleaved tree species and Picea abies (L.) Karst. wereplanted under spruce plantations of varying densities, in Sweden.Treatments included control (994 stems ha^–1), dense (538stems ha^–1), sparse (294 stems ha^–1) and gap (0stems ha^–1) overstory treatments. There was an increasein height and diameter growth from control to sparse overstorytreatment of all underplanted tree species except for ash (Fraxinusexcelsior L.) and Norway maple (Acer platanoides L.). Site conditionsmay have hampered the growth of these species, as well as wildcherry (Prunus avium L.). Both oak (Quercus robur L.) and sprucehad greater growth in the gap treatment, relative to the othertreatments. Insecticide application did not influence seedlinggrowth or survival. The performance of beech (Fagus sylvaticaL.), lime (Tilia cordata Mill.), spruce and oak was consistentwith shade tolerance ranking. Beech and lime had a very highsurvival rate, even under the densest canopy. The growth andmortality of ash, maple and wild cherry differed significantlyfrom what was expected. This experiment demonstrated significantvariation in interspecific growth and mortality between sevenplanted tree species in relation to canopy density. Correctsite and species selection is crucial when underplanting inshelterwood systems.     

Effect of tree species on carbon stocks in forest floor and mineral soil and implications for soil carbon inventories

Forest soil organic carbon (SOC) and forest floor carbon (FFC) stocks are highly variable. The sampling effort required to assess SOC and FFC stocks is therefore large, resulting in limited sampling and poor estimates of the size, spatial distribution, and changes in SOC and FFC stocks in many countries. Forest SOC and FFC stocks are influenced by tree species. Therefore, quantification of the effect of tree species on carbon stocks combined with spatial information on tree species distribution could improve insight into the spatial distribution of forest carbon stocks.We present a study on the effect of tree species on FFC and SOC stock for a forest in the Netherlands and evaluate how this information could be used for inventory improvement. We assessed FFC and SOC stocks in stands of beech (Fagus sylvatica), Douglas fir (Pseudotsuga menziesii), Scots pine (Pinus sylvestris), oak (Quercus robur) and larch (Larix kaempferi).FFC and SOC stocks differed between a number of species. FFC stocks varied between 11.1 Mg C ha⁻¹ (beech) and 29.6 Mg C ha⁻¹ (larch). SOC stocks varied between 53.3 Mg C ha⁻¹ (beech) and 97.1 Mg C ha⁻¹ (larch). At managed locations, carbon stocks were lower than at unmanaged locations. The Dutch carbon inventory currently overestimates FFC stocks. Differences in carbon stocks between conifer and broadleaf forests were significant enough to consider them relevant for the Dutch system for carbon inventory.     

Interspecific competition impacts on the morphology and distribution of fine roots in European beech (Fagus sylvatica L.) and Norway spruce (Picea abies (L.) Karst.)

Morphology and vertical distribution patterns of spruce and beech live fine roots (diameter ≤2 mm) were studied using a soil core method in three comparable mature stands in the Solling: (1) pure beech, (2) pure spruce and (3) mixed spruce–beech. This study was aimed at determining the effects of interspecific competition on fine root structure and spatial fine root distribution of both species. A vertical stratification of beech and spruce fine root systems was found in the mixed stand due to a shift in beech fine roots from upper to lower soil layers. Moreover, compared to pure beech, a significantly higher specific root length (SRL, P<0.05) and specific surface area (SSA, P<0.05) were found for beech admixed with spruce (pure beech/mixed beech SRL 16.1–23.4 m g⁻¹, SSA 286–367 cm² g⁻¹). Both indicate a flexible ‘foraging’ strategy of beech tending to increase soil exploitation and space sequestration efficiency in soil layers less occupied by competitors. Spruce, in contrast, followed a more conservative strategy keeping the shallow vertical rooting and the root morphology quite constant in both pure and mixed stands (pure spruce/mixed spruce SRL 9.6/7.7 m g⁻¹, P>0.10; SSA 225/212 cm² g⁻¹, P>0.10). Symmetric competition belowground between mixed beech and spruce was observed since live fine roots of both species were under-represented compared to pure stand. However, the higher space sequestration efficiency suggests a higher competitive ability of beech belowground.     

Effects of environmental changes on the vitality of forest stands

Using the physiological single tree growth model BALANCE, vitality of forest stands was simulated in dependence of the site-related factors, climate and stand structure. At six level II plots in southern Germany with the main tree species beech (Fagus sylvatica L.), oak (Quercus robur L.), spruce (Picea abies [L.] Karst.), and pine (Pinus sylvestris L.), simulated results were compared to measured values (soil water content, bud burst and leaf colouring, diameter at breast height, tree height and crown density) in order to validate the model. Sensitivity tests were done to examine the influence and the interactions of the environmental parameters. The validation results show that BALANCE is capable of realistically simulating the growth and vitality of forest stands for central European regions for medium term time spans (several years). The validation of the water balance module produces mean absolute errors based on field capacity between 2.7 and 6.9% in dependence of sites and forest stands. Senescence of foliage as well as crown density is reproduced with a correlation coefficient of 0.70 compared to measurements. Differences between measured and simulated diameter values were smaller than 1% for spruce and smaller than 6.5% for beech after 7 years of simulation, and smaller than 1% for oak after 8 years of simulation. On the other hand, the simulations for pine trees conform less with the measurements (difference: 22.6% after 8 years). The sensitivity of the model on environmental changes and on combinations of these parameters could be demonstrated. The responses of the forest stands were quite different.     

Regeneration growth in different light environments of mixed species, multiaged, mountainous forests of Romania

Growth of regenerating trees in different light environments was studied for the mountainous, mixed-species forests in the Carpathian Mountains of Romania. The primary species in these mixtures were silver fir (Abies alba Mill.), European beech (Fagus sylvatica L.) and Norway spruce (Picea abies (L.) Karst). Seedlings/saplings of these species were selected and measured in different stands from two different geographical locations. Regenerating trees were measured for height and diameter growth during the summer of 2002. For each seedling/sapling, percentage of above canopy light (PACL) and stand basal area (BA) were used to assess available and occupied growing space respectively. Regeneration growth was compared against these two variables and regression relationships were developed. Using these models, we predicted the dynamics of regeneration as both growth and species composition. Our results showed that in low-light environments (PACL<20–35%; BA>30 m²/ha), shade tolerant fir and beech clearly outcompeted the spruce. Therefore, in dense stands, spruce could be eliminated by the shade tolerant species. For intermediate levels of cover (PACL=35–70%; BA=15–35 m²/ha) the spruce grew at comparable rates as the beech and fir. All three species showed similar growth rates in open conditions (PACL>80–90%; BA<15–20 m²/ha) with the spruce having a tendency to outgrow the others. However, in terms of establishment, such conditions favor spruce and inhibit fir and beech.     

Comparison of soil nitrogen dynamics under beech,Norway spruce and Scots pine in central Germany

To investigate the effect of tree species on soil N dynamics in temperate forest ecosystems, total N (Nt), microbial N (Nmic), net N mineralization, net nitrification, and other soil chemical properties were comparatively examined in beech (64–68 years old) and Norway spruce (53–55 years old) on sites 1 and 2, and beech and Scots pine (45 years old) on site 3. The initial soil conditions of the two corresponding stands at each site were similar; soil types were dystric Planosol (site 1), stagnic Gleysols (site 2), and Podzols (site 3). In organic layers (LOf₁, Of₂, Oh), Nmic and Nmic/Nt, averaged over three sampling times (Aug., Nov., Apr.), were higher under the beech stands than under the corresponding coniferous ones. However, the Nmic in the organic layers under beech had a greater temporal variation. Incubation (10 weeks, 22 °C, samples from November) results showed that the net N mineralization rates in organic layers were relatively high with values of 8.1 to 24.8 mg N kg^–1 d^–1. Between the two corresponding stands, the differences in net N mineralization rates in most of the organic layers were very small. In contrast, initial net nitrification rates (0.2–17.1 mg N kg^–1 day^–1) were considerably lower in most of the organic layers under the conifer than under the beech. In the mineral soil (0–10 cm), Nmic values ranged from 4.1–72.7 mg kg^–1, following a clear sequence: August>November>April. Nmic values under the beech stands were significantly higher than those under the corresponding coniferous stands for samples from August and April, but not from November. The net N mineralization rates were very low in all the mineral soils studied (0.05–0.33 mg N kg^–1 day^–1), and no significant difference appeared between the two contrasting tree species.     

Effects of the extreme drought in 2003 on soil respiration in a mixed forest

We present a field study on the drought effects on total soil respiration (SR_t) and its components, i.e., “autotrophic” (SR_a: by roots/mycorrhizosphere) and “heterotrophic” respiration (SR_h: by microorganisms and soil fauna in bulk soil), in a mature European beech/Norway spruce forest. SR_a and SR_h were distinguished underneath groups of beech and spruce trees using the root exclusion method. Seasonal courses of SR_a and SR_h were studied from 2002 to 2004, with the summer of 2003 being extraordinarily warm and dry in Central Europe. We (1) analyzed the soil temperature (T _s) and moisture sensitivity of SR_a and SR_h underneath both tree species, and (2) examined whether drought caused differential decline of SR_a between spruce and beech. Throughout the study period, SR_a of beech accounted for 45–55% of SR_t, independent of the soil water regime; in contrast, SR_a was significantly reduced during drought in spruce, and amounted then to only 25% of SR_t. In parallel, fine-root production was decreased during 2003 by a factor of six in spruce (from 750 to 130 mg l⁻¹ a⁻¹), but remained at levels similar to those in 2002 in beech (about 470 mg l⁻¹ a⁻¹). This species-specific root response to drought was related to a stronger decline of SR_a in spruce (by about 70%) compared to beech (by about 50%). The sensitivity of SR_a and SR_h to changing T _s and available soil water was stronger in SR_a than SR_h in spruce, but not so in beech. It is concluded that SR_a determines the effect of prolonged drought on the C efflux from soil to a larger extent in spruce than beech, having potential implications for respective forest types. This article belongs to the special issue "Growth and defence of Norway spruce and European beech in pure and mixed stands."     

Does tree species composition control soil organic carbon pools in Mediterranean mountain forests?

We compared soil organic carbon (SOC) stocks and stability under two widely distributed tree species in the Mediterranean region: Scots pine (Pinus sylvestris L.) and Pyrenean oak (Quercus pyrenaica Willd.) at their ecotone. We hypothesised that soils under Scots pine store more SOC and that tree species composition controls the amount and biochemical composition of organic matter inputs, but does not influence physico-chemical stabilization of SOC. At three locations in Central Spain, we assessed SOC stocks in the forest floor and down to 50 cm in the mineral in pure and mixed stands of Pyrenean oak and Scots pine, as well as litterfall inputs over approximately 3 years at two sites. The relative SOC stability in the topsoil (0-10 cm) was determined through size-fractionation (53 μm) into mineral-associated and particulate organic matter and through KMnO₄-reactive C and soil C:N ratio.Scots pine soils stored 95-140 Mg ha⁻¹ of C (forest floor plus 50 cm mineral soil), roughly the double than Pyrenean oak soils (40-80 Mg ha⁻¹ of C), with stocks closely correlated to litterfall rates. Differences were most pronounced in the forest floor and uppermost 10 cm of the mineral soil, but remained evident in the deeper layers. Biochemical indicators of soil organic matter suggested that biochemical recalcitrance of soil organic matter was higher under pine than under oak, contributing as well to a greater SOC storage under pine. Differences in SOC stocks between tree species were mainly due to the particulate organic matter (not associated to mineral particles). Forest conversion from Pyrenean oak to Scots pine may contribute to enhance soil C sequestration, but only in form of mineral-unprotected soil organic matter.     

Carbon and nitrogen in forest floor and mineral soil under six common European tree species

The knowledge of tree species effects on soil C and N pools is scarce, particularly for European deciduous tree species. We studied forest floor and mineral soil carbon and nitrogen under six common European tree species in a common garden design replicated at six sites in Denmark. Three decades after planting the six tree species had different profiles in terms of litterfall, forest floor and mineral soil C and N attributes. Three groups were identified: (1) ash, maple and lime, (2) beech and oak, and (3) spruce. There were significant differences in forest floor and soil C and N contents and C/N ratios, also among the five deciduous tree species. The influence of tree species was most pronounced in the forest floor, where C and N contents increased in the order ash = lime = maple < oak = beech ? spruce. Tree species influenced mineral soil only in some of the sampled soil layers within 30 cm depth. Species with low forest floor C and N content had more C and N in the mineral soil. This opposite trend probably offset the differences in forest floor C and N with no significant difference between tree species in C and N contents of the whole soil profile. The effect of tree species on forest floor C and N content was primarily attributed to large differences in turnover rates as indicated by fractional annual loss of forest floor C and N. The C/N ratio of foliar litterfall was a good indicator of forest floor C and N contents, fractional annual loss of forest floor C and N, and mineral soil N status. Forest floor and litterfall C/N ratios were not related, whereas the C/N ratio of mineral soil (0–30 cm) better indicated N status under deciduous species on rich soil. The results suggest that European deciduous tree species differ in C and N sequestration rates within forest floor and mineral soil, respectively, but there is little evidence of major differences in the combined forest floor and mineral soil after three decades.     

Species mixing effect on Norway spruce response to elevated CO2 and climatic variables: root and radial growth response

A 7-year study was conducted to examine the growth (diameter and root) response of Norway spruce (Picea abies (L.) Karst.) seedlings to elevated CO₂ (CO_2ELV, 770 μmol (CO₂) mol^?1) in different mixture types (monospecific (M): a Norway spruce seedling surrounded by six spruce seedlings, group-admixture (G): a spruce seedling surrounded by three spruce and three European beech seedlings, single-admixture (S): a spruce seedling surrounded by six beech seedlings). After seven years of treatments, no significant effect from elevated CO₂ was found on the root dry mass (p?=?0.90) and radial growth (p?=?0.98) of Norway spruce. Neither did we find a significant interaction between [CO₂]?×?mixing treatments (p?=?0.56), i.e. there was not a significant effect of CO₂ concentrations [CO₂] in all the admixture types. On the contrary, spruce responses to admixture treatments were significant under CO_2AMB (p?=?0.05), which demonstrated that spruce mainly increased its growth (diameter and root) in M and neighbouring with beech was not favourable for spruce seedlings. In particular, spruce growth diminished when growing beside high proportions/numbers of European beech (S). Here, we also evaluated the association between tree-ring formation and climatic variables (precipitation and air temperature) in different admixture types under elevated and ambient CO₂ (CO_2AMB, 385 μmol (CO₂) mol^?1). Overall, our result suggests that spruce responses to climate factors can be affected by tree species mixing and CO₂ concentrations, i.e. the interaction between climatic variables?×?admixture types?×?[CO₂] could alter the response of spruce to climatic variables.

     

Carbon stocks and net ecosystem production changes with time in two Italian forest chronosequences

Forest management influences several ecosystem processes, including carbon exchange between forest ecosystem and atmosphere. The aim of this paper was to study the carbon cycle over different age classes of two managed forests in the Italian Alps through direct measurements and modelling. For this purpose, ecosystem carbon dynamics of a beech forest (Fagus sylvatica L.) and of a spruce forest (Picea abies (L.) Karst.) were investigated using a chronosequence approach. In both forests, five forest development stages were identified (thicket, pole wood, young forest, mature forest and the regeneration phase) with an age spanning from 42 to 163?years for the beech forest and from 35 to 161?years for the spruce forest. Measured total ecosystem carbon stock increased up to 80–100?years, with a mean of 232?MgC?ha^?1 in the beech forest and of 299?MgC?ha^?1 in the spruce forest. Calculated net ecosystem production (NEP) was found to decrease linearly with age and had an average value of 2.2 and 4.4?MgC?ha^?1?year^?1 for beech and spruce forest, respectively. Model simulations reported an increase in NEP till 50–60?years followed by a decrease thereafter. The model also predicted a negative NEP for a short period (8–11?years) after the seed cut. Aboveground biomass was the main driver of carbon accumulation while soil carbon was not significantly influenced by both age and management system. Moreover, measured data and model showed that the applied shelterwood system allowed for a rapid recovery of the ecosystem after the disturbance (i.e. seed cut), bringing back forest to act as C sink in few years.     

The increasing scarcity of red oaks in Mississippi River floodplain forests: Influence of the residual overstory

Red oaks – cherrybark oak (Quercus pagoda Raf.), willow oak (Quercus phellos L.), water oak (Quercus nigra L.), and Nuttall oak (Quercus texana Buckley; aka: Quercus nuttallii Palmer) – are not regrowing in Mississippi Delta river floodplain forests in the southeastern United States in sufficient numbers to sustain the former species composition and timber and wildlife values. Even if vigorous red oak reproduction becomes established, partial harvesting that does not remove the taller trees will suppress understory red oak height growth more than it will suppress height growth of such other species as sugarberry (Celtis laevigata Willd.), American elm (Ulmus americana L.), cedar elm (Ulmus crassifolia Nutt.), swamp dogwood (Cornus foemina Mill.), green ash (Fraxinus pennsylvanica Marshall), and sweetgum (Liquidambar styraciflua L.). Consequently, the red oaks in these partially harvested stands become increasingly suppressed and probably die; and there is a shift in species composition to the other species. In addition to ensuring vigorous oak reproduction, silvicultural clearcutting or rapid removal of the residual trees following shelterwood or seed tree harvesting to provide full sunlight is needed to ensure red oaks become a dominant part of these future river floodplain stands.     

Effects of litter quality and parent material on organic matter characteristics and N-dynamics in Luxembourg beech and hornbeam forests

To test effects of litter quality and soil conditions on N-dynamics, we selected seven forests in Luxembourg dominated by beech (Fagus sylvatica, L.) and hornbeam (Carpinus betulus L.), and located on acid loam, decalcified marl or limestone, and measured organic matter characteristics, microbial C and N and net N-mineralization in a laboratory incubation experiment. Organic layer characteristics were significantly affected by species, with lower litter decay and higher accumulation under the less palatable beech, even on limestone. However, beech and hornbeam did not show any differences in N-cycling at all. Instead of species, N-cycling was affected by site conditions, albeit different than expected. Microbial N generally increased from acid loam to limestone, but acid loam showed higher net N-mineralization, especially in the organic layer. Also, acid loam showed high instead of low efficiency of N-mineralization per unit microbe, in both organic layer and mineral topsoil. In addition, acid loam showed net consumption of DOC instead of release in both soil layers, which suggests that not N, but C was a limiting factor to decomposition. In contrast, limestone showed low net N-mineralization in the organic layer, despite high mass and well-decomposed organic matter, and low efficiency of N-mineralization per unit microbe in both organic layer and mineral topsoil. DOC was net released instead of consumed, which supported that not C, but N was a limiting factor. The general lack of differences in net N-cycling between species, but relatively clear site effects, is discussed in relation to different microbial strategies. Acid soil may have high net N-release despite low biological activity, because N-requirements of fungi are also low, while in calcareous soil, high bacterial N-demand may counteract high gross N-release. Thus, species producing litter that decomposes rapidly may be planted to improve soil conditions and plant biodiversity, but litter quality effects on N-availability may be less important than soil conditions.     

Changes in soil organic carbon fractions in a tropical Acrisol as influenced by the addition of different residue materials

Residues of Leucaena (Leucaena leucocephala (L), Senna siamea (S) and maize stover (M) were tested to evaluate their effect on soil organic matter accumulation and composition under sub-humid tropical conditions. On an Imperata cylindrica (I) dominated grass fallow, a total amount of 30?Mg?ha^?1 DM were applied within 18?months. Two months after the last application, changes in the light and heavy soil organic carbon fraction (LF and HF) and in the total soil organic carbon content (LF?+?HF) in the topsoil were observed. All organic materials increased the proportion of the LF fraction in the soil significantly. The increase in HF was 39 to 51% of the increase in total organic carbon, depending on the source of the organic material. The potential of the tested organic materials to increase total soil organic carbon content (including all soil organic carbon fractions) was in the order L?>?S?>?M?>?I, whereas the order of increase of the HF fraction was L?=?S?>?I?>?M. Cation exchange capacity of the newly formed heavy soil organic carbon was highest with L and lowest with M. Ranking of the transformation efficiency of applied plant residues into the heavy soil organic carbon fraction was I?>?L?=?S?>?M. Transformation efficiency of the residues could neither be explained by lignin nor lignin/N ratio, but rather by extractable polyphenols (Folin?CDenis extraction). The results show that accumulation of the HF fraction in tropical soils is feasible through the application of large quantities of plant residues, but depends strongly on the composition of the applied materials.     

Effects of roe deer browsing and site preparation on performance of planted broadleaved and conifer seedlings when using temporary fences

Abstract

The effects of fencing and site preparation on performance of birch (Betula pendula Roth), oak (Quercus robur L.), pine (Pinus sylvestris L.) and spruce [Picea abies (L.) Karst.] seedlings were studied over four growing seasons in southern Sweden. The experiment was composed of four browsing treatments, from which roe deer (Capreolus capreolus L.) were excluded for 0, 12, 24 or 42 months, and four soil treatments. Natural browsing was combined with artificial browsing (clipping). It was demonstrated that browsing by roe deer could be isolated from browsing by moose (Alces alces L.) by selective fencing. Except for birch, browsing had little effect on seedling survival. Browsing reduced height growth by more than 100% for oak and pine, and more than 60% of pine seedling developed multiple stems. Except for oak, site preparation increased seedling survival. Inverting site preparation, in combination with or without fertilization, produced positive growth responses in seedlings, whereas patch scarification did not. There was no interaction between browsing and soil treatments on seedling performance. It was concluded that long-term protection against browsing by roe deer is needed for oak and pine, whereas short-term fences might be used for birch, and that spruce is relatively unaffected by browsing.     

Influence of slope position, stand type and rhododendron (Rhododendron ponticum) on litter decomposition rates of Oriental beech (Fagus orientalis Lipsky.) and spruce [Picea orientalis (L.) Link]

Oriental beech (Fagus orientalis Lipsky.) and Oriental spruce [Picea orientalis (L.) Link] are the two most common tree species in northeast Turkey. Their distribution, stand type and understorey species are known to be influenced by topographical landforms. However, little information is available as to how these changes affect litter decomposition rates of these two species. Here, we investigated the effects of slope positions (top 1,800 m, middle 1,500 m and bottom 1,200 m), stand type (pure and mixed stands) and purple-flowered rhododendron (Rhododendron ponticum) on litter decomposition rates of Oriental beech and spruce for 4 years using the litterbag technique in the field. Among these three factors, stand type had the strongest influence on litter decomposition (P < 0.001, F = 58.8), followed by rhododendron (P < 0.001, F = 46.8) and slope position (P < 0.05, F = 11.6). Litter decomposition was highest under mixed beech/spruce forest, followed by pure beech and spruce forest. Beech and spruce litter decomposed much faster in mixed bags (beech–spruce) than they did separately under each stand type. Purple-flowered rhododendron significantly reduced litter decomposition of Oriental beech and spruce. Beech and spruce litter decomposed much slower at top slope position than at either bottom or middle position. Differential litter decomposition of Oriental beech and spruce was mainly due to adverse conditions in spruce forest and the presence of rhododendron on the ground which was associated with lower soil pH. Higher elevations (top slope position) slowed down litter decomposition by changing environmental conditions, most probably by decreasing temperature as also other factors are different (pH, precipitation) and no detailed investigations were made to differentiate these factors. The adverse conditions for litter decomposition in spruce forest can be effectively counteracted by admixture of beech to spruce monoculture and by using the clear-cutting method for controlling rhododendron.