Thirteen-year monitoring of liming and PK fertilization effects on tree vitality in Norway spruce and European beech stands

The chemical fertility of the forest soils in the Belgian Ardenne is threatened by acidifying and eutrophying deposition and by the nutrient removal due to timber harvesting. Experiments were launched to evaluate the ability of liming and fertilization to improve foliar nutrition, maintain or restore crown condition and promote tree growth. In 1995, 10 liming and fertilization trials were installed in even-aged stands of European beech (5) and Norway spruce (5) distributed throughout the Ardenne. In each stand, two treatments were tested: liming with 3,000 kg ha⁻¹ of dolomitic limestone and liming plus fertilization with 0 to 800 kg ha⁻¹ of rock phosphate and 0 to 350 kg ha⁻¹ of K₂SO₄. Between 1995 and 2006, the foliar Ca and Mg status of spruce and beech trees improved in the limed stands, which limited significantly but did not prevent the decline in crown condition triggered by the summer drought in 2003. For spruce, liming also increased the increment in basal area. The additional fertilization increased the foliar nutrition in P but had no significant effects on soil chemistry and tree vitality.     

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.     

Influence of stand,site and meteorological variables on the maximum leaf area index of beech,oak and Scots pine

Different multiple linear regression models of maximum leaf area index (LAI_max) based on stand characteristics, site quality, meteorological variables and their combinations were constructed and cross-validated for three economically important tree species in Flanders, Belgium: European beech (Fagus sylvatica L.), Pedunculate oak (Quercus robur L.) and Scots pine (Pinus sylvestris L.). The models were successfully tested on similar datasets of experimental sites across Europe. For each species, ten homogeneous and mature stands were selected, covering the species’ entire stand productivity range based on an a priori site index classification. LAI_max was derived from measurements of leaf area index (LAI) made by means of hemispherical digital photography over the whole growing season (mid-April till end October 2008). Species-specific models of LAI_max for beech and oak were mostly driven by management practice affecting stand characteristics and tree growth. Tree density and dominant height were main predictors for beech, while stand age and tree-ring growth were important in the oak models. Scots pine models were more affected by site quality and meteorological variables. The beech meteorological model showed very good agreement with LAI at several European sites. Scots pine’s stand model predicted well LAI across Europe. Since the species-specific models did not share common predictors, generic models of LAI_max were developed for the 30 studied sites. Dominant height was found to be the best predictor in those generic models. As expected, they showed a lower predictive performance than species-specific ones.     

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 exceeding the critical loads for nitrogen alter nitrate leaching,the nutrient status of trees and their crown condition at Swiss Long-term Forest Ecosystem Research (LWF) sites?

Nitrogen (N) deposition exceeds the critical loads for this element in most parts of Switzerland apart from the Alps. At 17 sites (8 broadleaved stands, 8 coniferous stands, and 1 mixed stand) of the Swiss Long-term Forest Ecosystem Research network, we are investigating whether N deposition is associated with the N status of the forest ecosystems. N deposition, assessed from throughfall measurements, was related to the following indicators: (1) nitrate leaching below the rooting zone (measured on a subset of 9 sites); (2) the N nutrition of the forest stand based on foliar analyses (16 sites); and (3) crown defoliation, a non specific indicator of tree vitality (all 17 sites). Nitrate leaching ranging from about 2 to 16 kg N ha⁻¹ a⁻¹ was observed at sites subjected to moderate to high total N deposition (>10 kg ha⁻¹ a⁻¹). The C/N ratio of the soil organic layer, or, when it was not present, of the upper 5 cm of the mineral soil, together with the pool of organic carbon in the soil, played a critical role, as previous studies have also found. In addition, the humus type may need to be considered as well. For instance, little nitrate leaching (<2 kg N ha⁻¹ a⁻¹) was recorded at the Novaggio site, which is subjected to high total N deposition (>30 kg ha⁻¹ a⁻¹) but characterized by a C/N ratio of 24, large organic C stocks, and a moder humus type. Foliar N concentrations correlated with N deposition in both broadleaved and coniferous stands. In half of the coniferous stands, foliar N concentrations were in the deficiency range. Crown defoliation tended to be negatively correlated with N concentrations in the needles. In the majority of the broadleaved stands, foliar N concentrations were in the optimum nutritional range or, on one beech plot with high total N deposition (>25 kg ha⁻¹ a⁻¹), above the optimum values. There was no correlation between the crown defoliation of broadleaved trees and foliar concentrations.     

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.     

Phosphorus uptake and maize response to organic and inorganic fertilizer inputs in Rubona,Southern Province of Rwanda

Green manure of multipurpose trees is known to be a good source of nutrients to crop. However, most agroforestry species do not have adequate phosphorus (P) in their leaves. Supplementing green manure with moderate dose of P is a beneficial strategy to improve food security in Rwanda. This study examines the effects of Calliandra calothyrsus Meissner, Tithonia diversifolia Hensley A.Gray and Tephrosia vogelii Hook.f. green manure applied independently or in combination with triple super phosphate (TSP) and lime on maize yield and P uptake in the Oxic Tropudalf of Rubona, Rwanda. The treatments were the control, lime at 2.5 t ha⁻¹, TSP at 25 and 50 kg P ha⁻¹, leaf of C. calothyrsus, T. diversifolia, and T. vogelii each at 25 and 50 kg P ha⁻¹, respectively. Leaf shrubs biomass, TSP and lime were applied for four consecutive seasons (2001–2004). The results showed that the combination of green manure with TSP at a rate of 50 kg P ha⁻¹ significantly increased maize yield from 24 to 508% when compared to the control and T. divesifolia combined with TSP was leading (508%). Equally, the same treatments as indicated above showed higher P uptake (15.6–18. 6 kg P ha⁻¹) than the control (5 kg P ha⁻¹) and 65% of maize yields variation was explained by total P uptake. The plant residues quality such as C:N ratio, total plant N, and P significantly influenced the variability of maize grain yields.     

Oak (<Emphasis Type="Italic">Quercus robur</Emphasis> L.) regeneration as a response to natural dynamics of stands in European hemiboreal zone

The oak (Quercus robur L.) regeneration intensity was assessed in the core area of the Białowieża National Park (BNP) in Poland with respect to the selected ecological factors. The emphasis was placed on the response of oak regeneration to disturbances, including the large-scale dieback of spruce stands. Defining their effect could help predicting the role of oak in naturally developing lowland forest ecosystems in the European hemiboreal zone. The results of the study challenge the opinion that the ‘lime-oak-hornbeam forest’ is a ‘climax’ community, confirming a very poor regeneration represented by only two saplings taller than 0.5 m per hectare. By contrast, in spruce-dominated communities, from 49 to 848 taller saplings per hectare were found. The occurrence of saplings was associated with discontinuous canopy of late seral stage of stands, as well as with large gaps. Most of the best quality grown-up oak saplings developed in the immediate neighbourhood of spruce logs. The results of the research indicate that ‘lime-oak-hornbeam forest’ (Tilio-Carpinetum) should be rather perceived as a transient community, evolved from relict, culturally modified, oak woodlands. Because spruce had become a dominating species only after abandoning in mid-1800s the historical regime of anthropogenic disturbances (involving frequent forest fires), the observed phenomena related to the disintegration of spruce stands had probably no precedent over the last 500 years. To confirm whether the massive decline of spruce stands will finally result in the successful establishment of the new canopy oak generation, both in Białowieża and other forests of hemiboreal zone, further research is needed.     

Stand density and growth of Norway spruce (<Emphasis Type="Italic">Picea abies</Emphasis> (L.) Karst.) and European beech (<Emphasis Type="Italic">Fagus sylvatica</Emphasis> L.): evidence from long-term experimental plots

On the basis of nine Norway spruce (Picea abies (L.) Karst.) and ten European beech (Fagus sylvatica L.) thinning experiments in Germany, for which both residual and removed stock had been registered first during 1870, I scrutinize how moderate and heavy thinning from below (B-, C-grade) affects the production of merchantable volume compared with light thinning (A-grade). In relation to A-grade, cumulative merchantable volume (CV) of B- and C-grade amounts in average to 103–107% in juvenile and to 97–102% in mature Norway spruce stands. The corresponding findings for European beech are 101–106% and 94–102%. CV of individual stands varies between 89% and 130% for Norway spruce and 73% and 155% for European beech (CV of A-grade = 100%). These findings are substantiated by the relation between stand density (SDI) and periodic annual increment (PAI). On the B- and C-grade plots of spruce and beech, respectively, SDI was reduced down to 41–91% and 31–83% of the A-grade. When SDI is reduced in young stands, PAI follows a unimodal curve. Norway spruce’s PAI culminates in 109% if SDI is reduced to 59%; European beech’s PAI culminates in 123% when density is reduced to 50%. Whereas Norway spruce’s growth reacts most positively on thinning under poor site conditions and with increment reduction on favourable sites, European beech behaves oppositely. With stand development the culmination point of the unimodal relation moves towards maximum density, so that in older stands PAI follows the increasing pattern, which is the left portion of a unimodal curve. A model is presented which apparently unifies contradictory patterns of stand density–growth reactions by integrating relative stand density, average tree size and site fertility effects, and makes the findings operable for forest management.     

Nutrient cycling and soil leaching in eighteen pure and mixed stands of beech (Fagus sylvatica) and spruce (Picea abies)

Studies on the combined effects of beech–spruce mixtures are very rare. Hence, forest nutrition (soil, foliage) and nutrient fluxes via throughfall and soil solution were measured in adjacent stands of pure spruce, mixed spruce–beech and pure beech on three nutrient rich sites (Flysch) and three nutrient poor sites (Molasse) over a 2-year period. At low deposition rates (highest throughfall fluxes: 17 kg N ha⁻¹ year⁻¹ and 5 kg S ha⁻¹ year⁻¹) there was hardly any linkage between nutrient inputs and outputs. Element outputs were rather driven by internal N (mineralization, nitrification) and S (net mineralization of organic S compounds, desorption of historically deposited S) sources. Nitrate and sulfate seepage losses of spruce–beech mixtures were higher than expected from the corresponding single-species stands due to an unfavorable combination of spruce-similar soil solution concentrations coupled with beech-similar water fluxes on Flysch, while most processes on Molasse showed linear responses. Our data show that nutrient leaching through the soil is not simply a “wash through” but is mediated by a complex set of reactions within the plant–soil system.     

Vulnerability of spruce (Picea abies) and beech (Fagus sylvatica) forest stands to storms and consequences for silviculture

The form and magnitude of storm damage and stand disclosure patterns were assessed in 332 randomly chosen pure and regular stands of spruce (Picea abies L.) and beech (Fagus sylvatica L.) after storm lothar, within a region of the Swiss Midlands. This data was analysed in relation to maximal wind speed, measured with Doppler radar techniques and other influential factors such as relief, allometric characteristics, silvicultural history, and neighbourhood. In addition, storm damage, assessed from aerial photographs over an extended perimeter (about 70,000 ha) was considered. A storm of the magnitude of lothar (December 26 1999), with an average maximal wind speed of 45 m s⁻¹ (160 km h⁻¹) appears to have a highly chaotic wind field structure, with great spatial and temporal variation of wind gusts. Wind speeds were not a significant predictor for damage in spruce stands and only weakly influential for beech. The consequences of this high randomness were analysed to estimate the return time of such a storm at the stand level. It lies between 86 and 113 years for spruce, 357 and 408 for beech. Only a few independent variables were significant and the overall explanatory strength of the model was unexpectedly low (R ²=0.07 for spruce and 0.30 for beech). Among the more reliable predisposing factors were mixture and aspect combined with gradient. An admixture of 10% or more broadleaved tree species or wind-firm conifers like Douglas fir [Pseudotsuga menziesii (Mirb.) Franco] significantly reduced the vulnerability of spruce stands (by a factor of more than three). On wind-exposed aspects, damage was more than twice the average. Steeper slopes caused a significant reduction in susceptibility (by a factor of six for slopes over 50%, in comparison to gentle slopes <20%). Other factors such as height to diameter ratio of trees or time since last thinning did not appear to be significant predictors.     

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.     

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.     

Organic amendment and inorganic fertilization affect soil properties and quality of Larix olgensis bareroot stock

Bareroot Changbai larch (Larix olgensis Henry.) seedlings were reared with inorganic fertilizer (nitrogen (N):phosphorus (P) = 1:1, W/W) applied at a rate of 100 (F100) or 200 kg N ha⁻¹ (F200) with (+) or without (−) chicken manure as a soil amendment (O) in north-eastern China. An unfertilized control treatment was included. Inorganic and organic fertilizer treatments tended to increase soil ammonium, nitrate, available P, total P, organic carbon content and electrical conductivity, and biomass and N concentration in seedlings. Organic amendment improved first order lateral root number, tap root length, fine root morphology (length, surface area, volume) in seedlings, while the F100 treatment increased N accumulation in needles and stems compared to the F200 treatment, on average. Most fertilizer treatments tended to increase P content in combined stems and roots, but F200 − O and F100 + O treatments diluted P in needles. Organic amendment combined with inorganic fertilizer at a rate of 100 kg N and P ha⁻¹ is recommended to improve seedling growth and N reserves in woody tissues.     

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.     

A Comparison Between European Continental and British Provenances of Some British Native Trees: Growth, Survival and Stem Form

Data describing the growth, survival and stem form of: a) Europeancontinental, and b) British provenances of trees native to Britainwere collated from Forestry Commission records. The growth ratesof European continental provenances of Scots pine (Pinus sylvestris),silver birch (Betulo pendula), sessile oak (Quercus petraea)and common alder (Alnus glutinosa) were inferior to those ofBritish provenances in 90 per cent of cases. European provenancesof these species also showed inferior survival. The growth ratesof continental provenances of beech (Fagus sylvatica) were superiorto British provenances in about 50 per cent of cases, provenancesfrom northern France, Belgium and Holland being the fastestgrowing. Relationships between the relative height growth offoreign provenances and latitude were demonstrated for Scotspine, silver birch and beech (relative height growth = the heightgrowth of a foreign provenance expressed relative to that ofBritish provenances at the same site). The continental provenancesthat grew best originated at 0–4° south of the latitudesof the planting sites. Relative height growth decreased bothnorth and south of these latitudes. In the case of oak, relativeheight growth did not vary widely between origins. The survivalof continental provenances of Scots pine was particularly pooron exposed upland sites. The stem form of British provenancesof Scots pine was intermediate between Fennoscandian provenancesand provenances from the rest of Europe. In oak and beech therewas generally little difference in stem form between Britishand continental provenances, but the best continental provenances(Belgian beech; French oak) were most reliable. The growth andsurvival of native trees is interpreted in the light of thepost-glacial history of these species. The implications regardingthe continued use of seed of native hardwoods imported fromcontinental Europe are discussed.     

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.     

Nutrient contents and efficiencies of beech and spruce saplings as influenced by competition and O<Subscript>3</Subscript>/CO<Subscript>2</Subscript> regime

Saplings of Fagus sylvatica and Picea abies were grown under conditions of intra and interspecific competition in a 2-year phytotron study under combinations of ambient and elevated ozone (+O₃ which is 2 × O₃, but <150 nl l⁻¹) as well as carbon dioxide concentrations (+CO₂ which is amb. CO₂ + 300 μl CO₂ l⁻¹) in a full factorial design. Saplings were analysed for various mineral nutrients in different plant organs as well as biomass production and crown development. The study was based on the assumption that nutritional parameters important for growth and competitiveness are affected by stress defence under limiting nutrient supply. The hypotheses tested were (1) that nutrient uptake-related parameters (a) as well as efficiencies in nutrient use for above-ground competition (b) of beech rather than spruce are impaired by the exposure to elevated O₃ concentrations, (2) that the efficiency in nutrient uptake of spruce is enhanced by elevated CO₂ concentrations in mixed culture, and (3) that the ability to occupy above-ground space at low nutrient cost is co-determinant for the competitive success in mixed culture. Clear nitrogen deficiencies were indicated for both species during the 2-year phytotron study, although foliar nitrogen-biomass relationships were not so close for spruce than for beech. O₃ stress did not impair nutrient uptake-related parameters of beech; thus hypothesis (1a). was not supported. A negative effect of elevated O₃ (under amb. CO₂) on the N and P based efficiencies in above-ground space occupation (i.e. lower crown volume per unit of N or P invested in stems, limbs and foliage) of beech supported hypothesis (1b). It appeared that ozone stress triggered a nutrient demand for stress defence and tolerance at the expense of above-ground competition (trade-off). Crown volume of beech under O₃ stress was stabilized in monoculture by increased nutrient uptake. In general, the +CO₂-treatment was able to counteract the impacts of 2 × O₃. Elevated CO₂ caused lower N and S concentrations in current-year foliage of both tree species, slightly higher macronutrient amounts in the root biomass of spruce, but did not increase the efficiencies in nutrient uptake of spruce in mixed culture. Therefore hypothesis (2) was not supported. At the end of the experiment spruce turned out to be the stronger competitor in mixed culture as displayed by its higher total shoot biomass and crown volume. The amounts of macronutrients in the above-ground biomass of spruce individuals in mixed culture distinctly exceeded those of beech, which had been strongly reduced by interspecific competition. The superior competitiveness of spruce was related to higher N and P-based efficiencies in above-ground space occupation as suggested in hypothesis (3). This article belongs to the special issue “Growth and defence of Norway spruce and European beech in pure and mixed stands”.     

Productivity of mixed versus pure stands of oak (Quercus petraea (Matt.) Liebl. and Quercus robur L.) and European beech (Fagus sylvatica L.) along an ecological gradient

The mixture of beech (Fagus sylvatica L.) and oak (sessile oak, Quercus petraea (Matt.) Liebl., and pedunculate oak, Q. robur L.) is of considerable importance in Europe and will probably become even more important under climate change. Therefore, the performance of oak and beech in mixture was compared with the species’ growth in pure stands. Data from 37 long-term mixing experiments in Poland, Germany and Switzerland were pooled for analysis of mixing effects on stand productivity and possible interrelationships with mixing portions or site conditions. We found that on average, mixed stands of oak and beech exceeded biomass productivity in pure stands by 30 % or 1.7 t ha^?1 year^?1, as the growth of both species was benefitted by the mixture. However, that the interaction actually ranged from facilitation and overyielding on poor sites to underyielding on fertile sites triggered by competition. An empirically derived interaction model showed volume and dry mass growth changing in mixed stands from gains of 50 % to losses of 10 % depending on site conditions. It is concluded that the analysed mixture grows in accordance with the stress-gradient hypothesis and that our results suggest a site-specific relationship between species mixture and biomass productivity. As a consequence, an adequate species mix should result in increased productivity under steady state as well as climate change.