Photosynthetic characteristics of Fagus sylvatica and Quercus robur established for stand conversion from Picea abies

Efforts in Europe to convert Norway spruce (Picea abies) plantations to broadleaf or mixed broadleaf-conifer forests could be bolstered by an increased understanding of how artificial regeneration acclimates and functions under a range of Norway spruce stand conditions. We studied foliage characteristics and leaf-level photosynthesis on 7-year-old European beech (Fagus sylvatica) and pedunculate oak (Quercus robur) regeneration established in open patches and shelterwoods of a partially harvested Norway spruce plantation in southwestern Sweden. Both species exhibited morphological plasticity at the leaf level by developing leaf blades in patches with an average mass per unit area (LMA) 54% greater than of those in shelterwoods, and at the plant level by maintaining a leaf area ratio (LAR) in shelterwoods that was 78% greater than in patches. However, we observed interspecific differences in photosynthetic capacity relative to spruce canopy openness. Photosynthetic capacity (A₁₆₀₀, net photosynthesis at a photosynthetic photon flux density of 1600 μmol photons m⁻² s⁻¹) of beech in respect to the canopy gradient was best related to leaf mass, and declined substantially with increasing canopy openness primarily because leaf nitrogen (N) in this species decreased about 0.9 mg g⁻¹ with each 10% rise in canopy openness. In contrast, A₁₆₀₀ of oak showed a weak response to mass-based N, and furthermore the percentage of N remained constant in oak leaf tissues across the canopy gradient. Therefore, oak photosynthetic capacity along the canopy gradient was best related to leaf area, and increased as the spruce canopy thinned primarily because LMA rose 8.6 g m⁻² for each 10% increase in canopy openness. These findings support the premise that spruce stand structure regulates photosynthetic capacity of beech through processes that determine N status of this species; leaf N (mass basis) was greatest under relatively closed spruce canopies where leaves apparently acclimate by enhancing light harvesting mechanisms. Spruce stand structure regulates photosynthetic capacity of oak through processes that control LMA; LMA was greatest under open spruce canopies of high light availability where leaves apparently acclimate by enhancing CO₂ fixation mechanisms.     

Differences in soil properties in adjacent stands of Scots pine, Norway spruce and silver birch in SW Sweden

Soil properties were compared in adjacent 50-year-old Norway spruce, Scots pine and silver birch stands growing on similar soils in south-west Sweden. The effects of tree species were most apparent in the humus layer and decreased with soil depth. At 20-30 cm depth in the mineral soil, species differences in soil properties were small and mostly not significant. Soil C, N, K, Ca, Mg, and Na content, pH, base saturation and fine root biomass all significantly differed between humus layers of different species. Since the climate, parent material, land use history and soil type were similar, the differences can be ascribed to tree species. Spruce stands had the largest amounts of carbon stored down to 30 cm depth in mineral soil (7.3 kg C m⁻²), whereas birch stands, with the lowest production, smallest amount of litterfall and lowest C:N ratio in litter and humus, had the smallest carbon pool (4.1 kg C m⁻²), with pine intermediate (4.9 kg C m⁻²). Similarly, soil nitrogen pools amounted to 349, 269, and 240 g N m⁻² for spruce, pine, and birch stands, respectively. The humus layer in birch stands was thin and mixed with mineral soil, and soil pH was highest in the birch stands. Spruce had the thickest humus layer with the lowest pH.     

Logging residue removal after thinning in Nordic boreal forests: Long-term impact on tree growth

The aim of this study was to determine the effect of whole-tree harvesting (WTH) on the growth of Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies (L.) Karst.) as compared to conventional stem harvesting (CH) over 10 and 20 years. Compensatory (WTH + CoF) and normal nitrogen-based (CH + F or WTH + F) fertilisation were also studied. A series of 22 field experiments were established during 1977-1987, representing a range of site types and climatic conditions in Finland, Norway and Sweden. The treatments were performed at the time of establishment and were repeated after 10-13 years at 11 experimental sites. Seven experiments were followed for 25 years.Volume increment was on average significantly lower after WTH than after CH in both 10-year periods in the spruce stands. In the pine stands thinned only once, the WTH induced growth reduction was significant during the second 10-year period, indicating a long-term response.Volume increment of pine stands was 4 and 8% and that of spruce stands 5 and 13% lower on the WTH plots than on CH during the first and the second 10-year period, respectively. For the second 10-year period the relative volume increment of the whole-tree harvested plots tended to be negatively correlated with the amount of logging residue. Accordingly, the relative volume increment decreased more, the more logging residue was harvested, stressing the importance of developing methods for leaving the nutrient-rich needles on site.If nutrient (N, P, K) losses with the removed logging residues were compensated with fertiliser (WTH + CoF), the volume increment was equal to that in the CH plots. Nitrogen (150-180 kg ha⁻¹) or N + P fertilisation increased tree growth in all experiments except in one very productive spruce stand. Pine stands fertilised only once had a normal positive growth response during the first 10-year period, on average 13 m³ ha⁻¹, followed by a negative response of 5 m³ ha⁻¹ during the second 10-year period. The fertilisation effect of WTH + F and WTH + CoF on basal area increment was both smaller and shorter than with CH + F.     

Dissolved organic carbon and nitrogen leaching from Scots pine, Norway spruce and silver birch stands in southern Sweden

The effects of three common tree species - Scots pine, Norway spruce and silver birch - on leaching of dissolved organic carbon and dissolved nitrogen were studied in an experimental forest with podzolised soils in southern Sweden. We analyzed soil water collected with lysimeters and modeled water fluxes to estimate dissolved C and N fluxes. Specific UV absorbance (SUVA) was analyzed to get information about the quality of dissolved organic matter leached from the different stands. Under the O horizon, DOC concentrations and fluxes in the birch stands were lower than in the spruce and pine stands; annual fluxes were 21 g m⁻² y⁻¹ for birch and 38 g m⁻² y⁻¹ and 37 g C m⁻² y⁻¹ for spruce and pine, respectively. Under the B horizon, annual fluxes for all tree species ranged between 3 and 5 g C m⁻² y⁻¹, implying greater loss of DOC in the mineral soil in the coniferous stands than in the birch stands. We did not find any effect of tree species on the quality of the dissolved organic matter, as measured by SUVA, indicating that the chemical composition of the organic matter was similar in leachates from all three tree species. Substantial amounts of nitrogen was leached out of the soil profile at the bottom of the B horizon from the pine and birch stands, whereas the spruce stands seemed to retain most of the nitrogen in the soil. These differences in N leaching have implications for soil N budgets.     

Factors affecting nutrient concentration and stoichiometry of forest trees in Catalonia (NE Spain)

Although some studies have observed significant correlations between latitude and climate gradients and tree leaf nutrient concentration and stoichiometry, others have not. This study examined the nutrient concentrations of tree leaves in 3530 plots of the Catalonian Forest Inventory. Catalonia is a Mediterranean region located in NE Iberian Peninsula. It has a long land-use history and includes the large industrial-urban area of Barcelona but still contains a large forest area (42%). In the forests of Catalonia, leaf nutrient concentration increased and leaf C:nutrient ratios decreased from south to north, which paralleled the increase in MAP (mean annual precipitation) and the decrease in MAT (mean annual temperature), which was expected in a Mediterranean climate where the availability of water is the most limiting factor for plant nutrient uptake. In addition, the availability of water, which influences productivity, was associated with low leaf N:P content ratios, which is consistent with the Growth Rate Hypothesis. At a regional scale, the results support the Soil-Age Hypothesis because the youngest soils in the Pyrenees had the lowest leaf N:P ratios. Furthermore, the type of forest (evergreen, deciduous, or coniferous) explained some of the variation in leaf nutrient concentrations and stoichiometry. Nutrient concentrations were highest in deciduous trees and lowest in coniferous trees. Leaf nutrient concentrations and stoichiometry were mainly correlated with climate, but other factors such as the chemical properties of soil and rock, phylogenetics, and different ecological histories and anthropogenic factors such as pollution, had an effect.     

Stocks and decay dynamics of above- and belowground coarse woody debris in managed Sitka spruce forests in Ireland

Coarse woody debris (CWD) has become recognised as an important component of the carbon (C) pool in forest ecosystems. In Ireland, managed Sitka spruce (Picea sitchensis (Bong) Carr.) forests account for 52.3% of the total forest estate. To determine the stock and decay dynamics of above and belowground CWD, field surveys using fixed area sample plots, were conducted in six even-aged Sitka spruce stands, representing the young, intermediate and mature stages of a typical commercial rotation. The volume, mass, density loss and C:N ratio of all CWD types (logs, stumps, and coarse roots) were determined using a five-decay class (DC) system. The decay rates and half life of CWD was also determined. To estimate CWD coarse root mass; roots associated with stumps classified in different decay classes were excavated. The coarse roots were categorised into small (2-10 mm), medium (10-50 mm) and large (>50 mm) diameter classes.CWD C-mass ranged from 6.98 to 18.62 Mg ha⁻¹ and was highest in an intermediate forest (D35), while the aboveground volume varied from 6.31 to 42.27 m³ ha⁻¹. Coarse roots accounted for 21% to 85% of the total CWD C-pool in the surveyed stands. The total CWD C-mass was poorly correlated with the number of thinning events (R² = 0.29), when data from D35 was excluded. The density loss was significant in logs (45%), stumps (58%), and small- (38%), medium- (50%) and large roots (38%) as decay progress from DC 0 to 4. There was a 46%, 41%, 51%, 72% and 57% decline in C:N ratio of logs, stumps, small-, medium- and large roots, respectively, as decay progressed from DC 0 to 4. The density decay rates were 0.059, 0.048 and 0.036 kg m⁻³ year⁻¹ for logs, stumps and coarse roots, respectively. The size classification of roots did not significantly affect their decay rate. The half life (50% decomposition) of CWD was estimated has 12-, 14- and 19 years for logs, stumps and roots of Sitka spruce. Regression curves showed a strong correlation between the density and C:N ratio (R² = 0.69, 0.74 and 0.93 for logs, stumps and coarse roots, respectively). The long term storage of C and its slow rate of decomposition make CWD a vital structural and functional component of the CWD C-pool and a major controller of forest ecosystem C-retention.     

Natural development and regeneration of a Central European montane spruce forest

Montane Norway spruce forests of Central Europe have a very long tradition of use for timber production; however, recently there has been increasing concern for their role in maintaining biological diversity. This concern, coupled with recent severe windstorms that led to wide-spread bark beetle outbreaks, has brought the management of montane spruce forests to the forefront of public policy discussions in Central Europe. In order to shed light on the natural development and current structure of mature montane spruce forests, we established four 0.25 ha research plots in a semi-natural montane spruce forest in the Šumava Mountains (The Bohemian Forest), Czech Republic. We mapped all trees, extracted increment cores for age and growth-pattern analyses, and inventoried all current tree regeneration, including the substrates on which it was found. Stands were characterized by uni-modal tree diameter distributions and high basal areas (56.6 m² ha⁻¹ on average), indicating a natural transition from the stem exclusion phase towards the understory reinitiation phase. The stands showed largely single-cohort recruitment age structures, however, with recruitment spanning seven decades. Our analyses suggest that this cohort existed as advance regeneration prior to major disturbances in the late 1800s, which included post-bark beetle salvage logging. Spatial pattern analyses of living and dead stems combined, showed an increase in uniformity of living trees, pointing to the role of natural density-dependent mortality. However, past growth patterns and historical documentation suggest that low intensity canopy disturbances (wind and snow) also caused mortality and diversified canopy structure. Because the stands developed naturally over the past 120+ years and thus escaped thinning operations, high volumes of coarse woody debris (94 m³ ha⁻¹) and snag densities (546 stems ha⁻¹) have accrued. Advance spruce regeneration was quite abundant and existed primarily on deadwood substrates, even though these occupied only a small percent of stand area. Because of salvage logging in the late 1880s, these stands do not qualify, according to the traditional paradigm, as natural spruce forests. As a result, they are recently subject to active management practices including salvage logging that remove dead and dying trees. Given the importance of deadwood for forest regeneration and recovery from disturbance, as demonstrated in this study, we argue that dead wood removal may limit future natural regeneration in these stands. Thus, the purported benefits of removing dead and dying trees from semi-natural forests must be carefully weighed against the potential detrimental impacts on natural spruce forest regeneration and biodiversity.     

Decomposing stumps influence carbon and nitrogen pools and fine-root distribution in soils

Decomposing stumps could significantly increase soil resource heterogeneity in forest ecosystems. However, the impact of these microsites on nutrient retention and cycling is relatively unknown. Stump soil was defined as the soil fraction directly altered by the decomposition of the primary rooting system (e.g. taproots) and aboveground stumps. Study sites were located in mature hardwood stands within the Jefferson National Forest in the Ridge and Valley Physiographic region of southwest Virginia. The objectives of this study were to: (i) determine the total soil volume altered by the decomposition of stumps and underlying root system, (ii) compare and contrast total C and N, extractable ammonium (NH₄⁺) and nitrate (NO₃⁻), potentially mineralizable N, microbial biomass C (MBC), root length and root surface area between the bulk soil (i.e. O, A, B and C horizons) and stump soil and (iii) evaluate how nutrient concentrations and fine-root dynamics change as stumps decompose over time using a categorical decay class system for stumps. Potentially mineralizable N was 2.5 times greater in stump soil than the A horizon (103 mg kg⁻¹ vs. 39 mg kg⁻¹), 2.7 times greater for extractable NH₄⁺ (16 mg kg⁻¹ vs. 6 mg kg⁻¹) and almost 4 times greater for MBC (1528 mg kg⁻¹ vs. 397 mg kg⁻¹). Approximately 19% of the total fine-root length and 14% of fine-root surface area occurred in the stump soil. Significant differences occurred in C and N concentrations between all four decay classes and the mineral soil. This validated the use of this system and the need to calculate weighted averages based on the frequency and soil volume influenced by each decay class. In this forest ecosystem, approximately 1.2% of the total soil volume was classified as stump soil and contained 10% and 4% of soil C and N. This study illustrates that including stump soil in soil nutrient budgets by decay class will increase the accuracy of ecosystem nutrient budgets.     

Dynamics of seedling establishment and survival in uneven-aged boreal forests

The aim of this study was to describe the dynamics of seedling establishment and development in spruce-dominated uneven-aged boreal forests. The study was based on empirical data from 15 stands with permanent plots, which had been intensively monitored for 10 years in southern Finland. All trees (height > 1.3 m) were measured every fifth year. Regeneration was measured on 64 permanent sample plots (4 m²) in each stand. The establishment of first-year seedlings was analysed on a yearly basis. The survival and development of older Norway spruce (Picea abies (L.) H. Karst.) seedlings were analysed based on observations made every five years. The establishment of spruce seedlings was closely correlated with the abundance of seed crops. Seedling cohorts originating from abundant seed crops were clearly detectable in the development of seedling height distributions over time. It took about 15 years for spruce seedlings to reach a height level of 15-30 cm. Local basal area had hardly any effect on the emergence or survival of small spruce seedlings, while the number of higher spruce seedlings decreased with increasing local basal area.     

The impact of atmospheric deposition and climate on forest growth in European monitoring plots: An individual tree growth model

In the climate change discussion, the possibility of carbon sequestration of forests plays an important role. Therefore, research on the effects of environmental changes on net primary productivity is interesting. In this study we investigated the influence of changing temperature, precipitation and deposition of sulphur and nitrogen compounds on forest growth. The database consisted of 654 plots of the European intensive monitoring program (Level II plots) with 5-year growth data for the period 1994–1999. Among these 654 plots only 382 plots in 18 European countries met the requirements necessary to be used in our analysis. Our analysis was done for common beech (Fagus sylvatica), oak (Quercus petraea and Q. robur), Scots pine (Pinus sylvestris) and Norway spruce (Picea abies). We developed an individual tree growth model with measured basal area increment of each individual tree as responding growth factor and tree size (diameter at breast height), tree competition (basal area of larger trees and stand density index), site factors (soil C/N ratio, temperature), and environmental factors (temperature change compared to long-term average, nitrogen and sulphur deposition) as influencing parameters. Using a mixed model approach, all models for the tree species show a high goodness of fit with Pseudo-R² between 0.33 and 0.44. Diameter at breast height and basal area of larger trees were highly influential variables in all models. Increasing temperature shows a positive effect on growth for all species except Norway spruce. Nitrogen deposition shows a positive impact on growth for all four species. This influence was significant with p < 0.05 for all species except common beech. For beech the effect was nearly significant (p = 0.077). An increase of 1 kg N ha⁻¹ yr⁻¹ corresponds to an increase in basal area increment between 1.20% and 1.49% depending on species. Considering an average total carbon uptake for European forests near 1730 kg per hectare and year, this implies an estimated sequestration of approximately 21–26 kg carbon per kg nitrogen deposition.     

Soil solution,foliar concentrations and tree growth response to 3-year of ammonium-nitrate addition in two boreal forests of Québec,Canada

Ammonium nitrate (NH₄NO₃) was applied monthly (from June to October) for 3 years in a balsam fir (Abies balsamea (Linné) Miller) and a black spruce (Picea mariana (Mill.) BSP) boreal forest in Québec (Canada). The design was composed of nine experimental units of 10 m × 10 m for each site. Application rates were 3 and 10 times the atmospheric N deposition measured at each site which was 6 and 3 kg ha⁻¹ year⁻¹ for the fir and the spruce sites, respectively. Soil solution composition (30 and 60 cm), tree growth, and foliar concentrations were analysed. The inorganic N in the soil solution of the control plots of both sites was low, particularly at the spruce site indicating that these forests are actively accumulating the atmospheric deposited N. Nitrogen additions regularly caused sudden and large inorganic N increases in the soil solution at both sites, both treatments and both sampling depths. However, these increases were transitory in nature and no persistent changes in inorganic N were observed. It was estimated that more than 95% of the added N was retained above the rooting zone at both sites. Nitrogen addition increased N, Ca, Mg and Mn foliar concentrations at the black spruce site but had no effects at the balsam fir site. After 3 years of N application, tree growth was similar in the control and the treated plots at both sites. Our results show that slow growing black spruce boreal forests with low ambient N deposition are responsive (in term of foliar N, Ca, Mg and Mn concentrations) to even small increases in N inputs, compared to higher growth balsam fir boreal forests with higher N deposition.     

Long-term dynamics of a mixed conifer stand in Slovenia managed with a farmer selection system

The single-tree selection system is an important option for management of Norway spruce (Picea abies (L.) Karst.) and silver fir (Abies alba Mill.) forests because it provides continuous cover, requires low investments for tending, and promotes natural regeneration as well as high stand resistance and elasticity. It is often regarded as a very conservative system that usually results in only minor spatiotemporal changes in forest structure and composition. We studied management history, structural changes, regeneration dynamics, and light climate of a traditional single-tree farmer selection silver fir-Norway spruce forest (site typology Bazzanio-Abietetum). Stand structure was analyzed on five 0.25 ha permanent plots in 1994, 2001, and 2008. Regeneration density and height growth, forest floor vegetation, and light climate were also assessed on 1.5 × 1.5 m regeneration subplots in 2001 and 2008. Tree cores extracted from dominant trees from both species in two plots were used for reconstructing stand history and age structure of the canopy layer. We documented the forest response to three types of selection management regimes: excessive, normal, and conservative. Excessive management with harvest intensity significantly above the increment was documented until the late 1950s, including two peaks of heavy fellings (diameter limit cut) in the 1880s and 1930s, which favoured establishment of Norway spruce and released regeneration. The period that followed was characterized by normal selection management, but was nevertheless marked by a decline of silver fir as a result of air pollution and several droughts. This led to sanitary fellings that were carried out from the late 1970s to the early 1990s. In the last two decades conservative management followed, which led to suppression and decline of regeneration, especially of Norway spruce, and loss of selection structure. Although we recorded lower regeneration potential of silver fir compared with Norway spruce within the seedling category, silver fir outcompeted Norway spruce within the small-sized tree category (1 cm < dbh ? 10 cm) because of its superior height growth in low light levels (diffuse light <6%) and occupied a greater share of the canopy. Nevertheless, we anticipate that over the long-term the low light regime will also cause regeneration decline of silver fir and broadleaves. Our research revealed significant structural changes in a single-tree farmer selection forest during the last 150 years. These were a result of variable management regime and environment. A farmer single-tree selection system could better mimic the natural disturbance regime if spatiotemporal combinations of diverse felling regimes would be used.     

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.     

Nutrient contents and concentrations in relation to growth of Picea abies and Fagus sylvatica along a European transect

Mineral nutrition of Norway spruce (Picea abies (L.) Karst.) and beech (Fagus sylvatica L.) was investigated along a transect extending from northern Sweden to central Italy. Nitrogen (N) concentrations of needles and leaves in stands growing on acid soils did not differ significantly between central Italy and southern Sweden (1.0 +/- 0.1 mmol N g(-1) for needles and 1.9 +/- 0.14 mmol N g(-1) for leaves). In both species, foliar N concentrations were highest in Germany (1.2 mmol N g(-1) for needles and 2.0 mmol N g(-1) for leaves) and decreased by 50% toward northern Sweden (0.5 mmol N g(-1)). Both species showed constant S/N and P/N ratios along the transect. Calcium, K and Mg concentrations generally reflected local soil conditions; however, Mg concentrations reached deficiency values in Germany. Leaf area per unit dry weight varied significantly along the transect with lowest values for Norway spruce recorded in northern Sweden and Italy (3.4 m(2) kg(-1)) and a maximum in central Europe (4.7 m(2) kg(-1)). A similar pattern was observed for beech. Despite the low variation in foliar N concentrations on the large geographic scale, local and regional variations in N concentrations equalled or exceeded the variation along the entire continental transect. Furthermore, nutrient contents (i.e., nutrient concentration x dry weight per needle or leaf) showed a greater variation than nutrient concentrations along the transect. Nitrogen contents of Norway spruce needles reached minimum values in northern Sweden (2.4 micro mol N needle(-1)) and maximum values in Denmark (5.0 micro mol N needle(-1)). The N content of beech leaves was highest in Denmark (242 micro mol N leaf(-1)). At the German site, foliar N content rather than N concentration reflected the seasonal dynamics of foliar growth and N storage of the two species. During foliage expansion, there was an initial rapid increase in N content and a decrease in N concentration. This pattern lasted for about 2 weeks after bud break and was followed by 6 weeks during which dry weight and N content of the foliage increased, resulting in a further decrease in N concentration. During summer, dry weight and N content of mature needles of Norway spruce increased further to reach a maximum in autumn, whereas N concentration remained constant. In spring, reallocation of N from 1- and 2-year-old needles was 1.5 and 1.0 micro mol N needle(-1), respectively. This remobilized N was a major source of N for the development of new needles, which had an N content of 1.5 micro mol N needle(-1) after bud break. The seasonal remobilization of N from old foliage decreased with increasing needle age. Needle N content and dry weight decreased progressively with age (1 micro mol N needle(-1) between age classes 2 and 5), whereas N concentrations remained constant. For Norway spruce, annual stemwood production was correlated with needle N content but not with foliar N concentration or with the total amount of N in the canopy. Interspecific and geographical differences in plant nutrition are discussed on the basis of competitive demands for C and N between growth of foliage and wood.     

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."     

Effects of wood-ash application on potential carbon and nitrogen mineralisation at two forest sites with different tree species,climate and N status

In the future it may become common practice to return wood-ash to forest ecosystems in order to replenish nutrients removed when brash has been extracted as a source of bioenergy. Wood-ash contains most of the nutrients that are present in the brash before its removal and burning, with the important exception of nitrogen (N). In the present paper we report measurements of CO₂ emissions and net N mineralisation in the humus layer and the upper 5 cm of mineral soil 12 years after the application of wood-ash to two study sites, representing different tree species, climatic conditions and N deposition histories. We hypothesized that application of wood-ash would increase both carbon (C) and net N mineralisation rates at Torup, an N-rich site with Norway spruce (Picea abies (L.) Karst.) in the south, whereas the net N mineralisation rates would not be affected at Vindeln, an N-poor site with Scots pine (Pinus sylvestris L.) in the north, where a possible N-limitation would restrict any N mineralisation. The treatments, comprising additions of 0, 1, 3 or 6 Mg of granulated wood-ash ha⁻¹, were applied in a randomised block design, replicated three times. Wood-ash from the same batch was used for all treatments at both sites. All factors were measured under laboratory conditions with controlled temperature and moisture levels. The potential CO₂ emissions (kg ha⁻¹ year⁻¹ of CO₂–C) at Torup were significantly higher in the 3 and 6 Mg ha⁻¹ treatments than in the control treatment, and the highest application resulted in an extra loss of 0.5 Mg ha⁻¹ of soil C annually as compared to the control. No such differences were detected at Vindeln. The results suggest that wood-ash application can deplete soil organic C at locations with similar characteristics (N-rich soil, spruce dominated, warm climate) as at Torup in this study.     

Sap flow of birch and Norway spruce during the European heat and drought in summer 2003

In this study the hydrological regime of Norway spruce (Picea abies) and birch (Betula pendula) growing on heavy soils in the south east of Austria was analysed. Results from the year 2003 characterised by an extremely hot and dry summer are presented in this paper. Due to the extreme weather conditions the soil water content in August 2003 was very low (0.10–0.25 m³ m⁻³) in the topsoil (0–50 cm) with no explicit difference between both tree species.     

Long term accumulation of nitrogen in soils of dry mixed eucalypt forest in the absence of fire

In the Eden area in NSW, Australia, low fertility granitic surface soils were sampled from 156 sites and analysed for pH, organic C, total N, total P, available P, exchangeable bases and exchangeable Al. Fifty eight of these sites were also sampled to a depth of 40 cm. Time since fire ranged from 1 to 39 years and was used in the analysis as a surrogate for fire frequency. No information was available on fire intensity. No significant relationships were found between time since fire and P or base cations. However, the quantities of organic matter and total N (kg ha⁻¹), and the C/N ratio were significantly related to both time since fire alone and to the combination of time since fire and soil total P. Based on these relationships, it was estimated that there were average net increases of between 11 and 21 kg N ha⁻¹ year⁻¹ in surface soil, the actual quantity depending on the level of soil total P. There was little change in N in the initial 10 years after fire and there was a peak in N accumulation about 24 years after fire. The C/N ratio and surface soil pH decreased with time since fire. Accumulation of N and reductions in pH and C/N ratio were studied further in a small scale paired plot analysis. The repeatedly burnt plots had lower levels of both litter and understorey and the overstorey trees generally had healthier crowns than in the unburnt plots. The differences between the repeatedly burnt and the unburnt plots matched the models developed from the general survey. There were no significant changes in the C/N ratio, but the unburnt sites had higher levels of extractable mineral N and the relationships between the mineral N and the C/N ratio for burnt and unburnt sites were statistically significant. The quantities of extractable mineral N in the unburnt soils (2.3 kg N ha⁻¹) were about twice the levels in the burnt soils (1.2 kg N ha⁻¹). The pH of the surface soil (4.4 in 1:1 water) in the regularly burnt area was higher than in the unburnt area (pH 4.1) and the exchangeable aluminium also differed (0.62 c mol⁻¹ in the burnt area and 1.3 c mol⁻¹ in the unburnt). The combined data indicate that changes occur in forest soils when there is a long period of exclusion of fire. It is suggested that these changes generally lead to secondary changes, such as in pH and availability of other elements such as aluminium. The study highlights a number of issues including the rates of inputs of N to the system and the question of N saturation and its long term interaction with plant species. It is hypothesised that reduced burning leads to increased N availability and other soil changes which negatively impact on tree health.     

Litterfall and nutrient return in five tree species in a common garden experiment

Canopy litterfall is a significant pathway for return of nutrients and carbon (C) to the soil in forest ecosystems. Litterfall was studied in five even-aged stands of Norway spruce, Sitka spruce, Douglas-fir, European beech and common oak at three different locations in Denmark; two sandy sites, Ulborg and Lindet in Jutland, and one loamy site, Frederiksborg on Zealand. Litterfall was collected during three years from 1994 to 1996 in all five species and during six years from 1994 to 1999 in Norway spruce, Sitka spruce and European beech. The average total litterfall was in the range of 3200–3700 kg ha⁻¹ yr⁻¹ and did not differ significantly among tree species. There were no significant differences in total litterfall among sites during the short period, but during the longer period the richer site Frederiksborg had significantly higher total and foliar litterfall amounts compared to the more nutrient-poor sites Lindet and Ulborg. There were close relationships between foliar and total litterfall suggesting that foliar litterfall can be reliably estimated from total litterfall. Beech and oak bud scale litter was significantly related to foliar litterfall. The amount of branch and twig litter was significantly higher in oak than in other tree species. The average foliar litterfall was well related to the annual volume increment. The relationship differed markedly from previously reported relationships based on global litterfall data suggesting that such relationships are better evaluated at the regional level. Nutrient concentrations and fluxes in foliar litterfall were not significantly different among the five tree species. However, there was a significant effect of site on most nutrient concentrations of the three litterfall fractions, and foliar fluxes of P, Ca and Mn were all significantly highest at Frederiksborg and lowest at Ulborg. The similarity in litterfall inputs to the forest floor under these five tree species suggested that previous reports of large variability in forest floor accumulation should primarily be attributed to differences in litter decomposition.     

Effects of experimental drought on the fine root system of mature Norway spruce

Norway spruce (Picea abies (L.) Karst.) is an economically important, but relatively drought-sensitive tree species that might suffer from increasing drought intensities and frequencies, which are predicted to occur in parts of central Europe under future climatic change. In a throughfall exclusion experiment using sub-canopy roofs, we tested the hypothesis that enhanced drought leads to an increase in fine root mortality, and also to a higher, subsequent fine root growth. Fine root production and mortality were assessed using two independent approaches, sequential soil coring (organic layer) and direct observations in minirhizotrons (organic layer plus upper mineral soil). Six weeks of throughfall exclusion resulted in mild drought stress, i.e. a reduction in average soil moisture from 20 to 12 vol.% during the treatment. Based on the sequential coring data, experimental drought did not result in significant changes in fine root biomass during the 6-week treatment period, but caused an increase in fine root mortality by 61% in the 6 weeks following the drought treatment. Remarkably, fine root production showed a synchronous increase in this period, which more than compensated for the loss due to increased mortality. The minirhizotron data confirmed that the drought treatment increased fine root loss in the organic layer. Based on this method, however, root loss occurred during the drought period and was not compensated by increased root production. The mild drought stress was mainly restricted to the organic layer and did not significantly influence fine root dynamics in the mineral soil. We calculated that the drought event resulted in an extra input of about 28 g C m⁻² and 1.1 g N m⁻² to the soil due to increased fine root mortality. We conclude that even periods of mild drought significantly increase fine root mortality and the associated input of root-derived C to the soil organic matter pool in temperate Norway spruce forests.