Pedunculate oak (Quercus robur L.) silviculture in natural stands of NW Spain: Environmental conditioners

Forest management is often carried out in different ways, without any appropriate environmental restrictions. Stands of pedunculate oak (Quercus robur L.) in Galicia (NW Spain) have been harvested by alternating high forest and, mainly, coppice forest. However, some totally inappropriate silvicultural treatments have been used, such as thinning of the best trees and inadequate pruning. The objective of the present study was to analyse how environmental characteristics affect the management of oak forests in Galicia. For this, a botanical inventory was carried out in 39 selected stands of Q. robur and a total of 42 parameters were measured, 4 of which were physiographical, 12 climatic, 19 edaphic and 7 silvicultural. In order to analyse the possible relationships among these variables, the silvicultural data were compared with the other data, by canonical correlation analysis. All parameters were correlated with the silvicultural regime, although the correlation was weak for the floristic data. It is therefore evident that the environmental conditions affect how forest stands should be managed, although this does not imply that more profitable use of the stands cannot be achieved than at present, and alternative silvicultural methods must be found to enable appropriate management and conservation of oak stands.     

Genetic structure of the green oak leaf roller (Tortrix viridana L.) and one of its hosts, Quercus robur L.

An herbivorous insect, the green oak leaf roller (Tortrix viridana) and one of its host species, pedunculate oak (Quercus robur), were investigated for their genetic variation within and among populations on the basis of mitochondrial, chloroplast, and anonymous markers. Oaks and green oak leaf rollers from 10 oak stands in North Rhine-Westphalia (North-Western Germany) were examined. The genetic variation of pedunculate oaks is much lower within than among populations when assessed using chloroplast gene markers and higher among populations when assessed using anonymous markers. When using mitochondrial and AFLP markers, the genetic variation of the green oak leaf roller populations was higher within than among the populations, which suggests a high gene flow between the populations. Mantel tests on the gene diversities of oaks and the green oak leaf roller yielded a significant negative correlation for both marker types. To sum up, the reasons for the differences in the spatial patterns of the genetic variation of the host and herbivorous insect may be found in their different generation times, mechanisms and capacities for dispersal.The results for the green oak leaf roller revealed a higher migration rate than assumed before which has consequences for the prediction of the dynamics of future outbreak events.     

Establishment and growth of oak (Quercus alba, Quercus prinus) seedlings in burned and fire-excluded upland forests on the Cumberland Plateau

Recurrent problems with regeneration of oaks (Quercus spp.) have been documented across a wide range of ecosystems. In oak-dominated forests of the central and Appalachian hardwood regions of the United States, a lack of competitive oak regeneration has been tied, in part, to fire suppression in these landscapes, and managers throughout the region are using prescribed fire to address this concern. To examine fire effects on oak regeneration, researchers have generally relied on inventories or population studies of existing seedlings. These studies are valuable but do not permit examination of the role of fire in enhancing the establishment and growth of new oak seedlings stemming from oak mast events. In this study, white (Quercus alba) and chestnut oak (Quercus prinus) acorn mast crops serendipitously occurred in year three (fall 2005) of a landscape-scale prescribed fire experiment. We examined establishment, survival, height and diameter of new seedlings on sites on the Cumberland Plateau in eastern Kentucky. Treatments were fire exclusion, a single prescribed fire (1x-burn; 2003), and repeated prescribed fire (3x-burn; 2003, 2004, and after acorn drop in 2006), all conducted in late spring. Initial densities of newly established chestnut and white oak seedlings were statistically similar across treatments (P = 0.42), despite fires on the 3x-burn site having occurred after acorns were on the ground. Oak seedling density was significantly predicted by oak basal area on all sites (R² = 0.12–0.46), except for chestnut oak on fire-excluded sites (R² = 0.04). Litter depth was less on 3x-burn sites compared to 1x-burn and fire-excluded sites, whereas canopy openness was greater on both burn treatments compared to fire-excluded sites. Seedling mortality was generally higher on fire-excluded sites compared to burn sites, especially for white oak. Oak seedling mortality in the first two growing seasons was significantly predicted by initial litter depth and open sky, with greater litter depth and lower percent open sky leading to higher mortality. In the third growing season none of the measured variables predicted chestnut oak seedling survival; for white oak, percent open sky remained a significant predictor of mortality. Initially, seedlings on the fire-excluded sites had similar height but smaller diameter; after three growing seasons there were few differences in seedling height or diameter among treatments. Our findings suggest a potential role for prescribed fire in establishing forest floor and light conditions that may enhance the success of new oak germinants, although different responses among species may suggest the need to target management for individual oak species.     

Wild boars (Sus scrofa) affect the recruitment rate and spatial distribution of holm oak (Quercus ilex)

The role of herbivores in controlling plant population abundance and distribution is unclear. We experimentally determine the effect of damage by wild boars (Sus scrofa) in recruitment rate and spatial pattern of a Mediterranean tree, the holm oak (Quercus ilex). We monitored oak establishment in the Sierra Nevada of southeastern Spain during 4 years (1999–2002) in four plots, two fenced and two unfenced that were used as controls. In addition, we planted 1-year-old seedlings in all microhabitats, both in fenced and control plots, to experimentally determine the effect of wild boar on their microhabitat-dependent survival. Despite seedling abundance being similar inside and outside the fences, sapling abundance diminished up to 50% in unfenced plots compared to fenced plots. Wild boars also seemed to modify the spatial distribution of seedlings and saplings. Whereas seedlings were most abundant under pines both in fenced and control plots, sapling abundance was highest under pine trees in fenced plots, but similarly abundant under pines or shrubs in control plots. This change in spatial pattern was due to the effect of wild boars killing seedlings when looking for food under pines. The effects of the different mortality factors were not additive. Where ungulates are absent, recruitment is high and occurs under oaks and especially under pines. Where wild boars are present, recruitment diminishes and occurs mostly under shrubs and pines. Wild boars arrest population growth and modify the spatial pattern of recruitment.     

Bacillus pumilus and Stenotrophomonas maltophilia as two potentially causative agents involved in Persian oak decline in Zagros forests (Iran)

The oak decline is known as one of the most destructive complex diseases causing high economic losses around the world, especially in Iran. The main objective of the present study was to investigate the possible role of bacteria as causative agents of oak decline in the Zagros forests of Iran. To do this, stem, root and leaf samples were taken from symptomatic Persian oak trees (Quercus brantii) in different zones of Zagros forests (Ilam Province, Iran). From 150 bacterial isolates, 20 showed pathogenicity against Geranium seedlings. Among 20 hypersensitivity test positive strains, four strains showed pathogenicity against oak saplings. Based on morphological and 16S rRNA gene sequence analysis, three strains were identified as Bacillus pumilus and one strain as non‐sporulating Gram‐negative Stenotrophomonas maltophilia. Pathogenicity studies of different B. pumilus and S. maltophilia strains revealed that they have potential to cause the disease in oak saplings and symptoms of disorder in Persian oak trees. To our knowledge, there are no previous records of B. pumilus and S. maltophilia causing decline on Fagaceous trees like Q. brantii. More detailed field and molecular studies are required to confirm the absolute role of such bacteria in occurrence of oak decline in Zagros forests.     

Modelling jay (Garrulus glandarius) abundance and distribution for oak regeneration assessment in Mediterranean landscapes

Because natural regeneration of oak is strongly dependent on jay abundance and distribution, we need to understand the determinants of jay abundance and occurrence as a first step to assess oak regeneration. In this paper we modelled the jay population distribution and abundance in a Mediterranean landscape mosaic of the eastern Iberian Peninsula (Valencia, east Spain). The methodology was based on assessing landscape attributes (habitat composition and configuration variables) on seven 9-km² sites and registering jay fledgling locations. Using a stepwise regression model we determined the variables that best explained the jay density. The probability of occurrence of jay nests within each site was assessed by comparing the landscape attributes of jay and non-jay areas within each site. Results were validated by predicting each site on the basis of data from the remaining six sites and then calculating the deviation between the predicted and the observed values in the field. The results suggest that jay density correlates positively with forest cover and landscape heterogeneity variables, and negatively with shrubland cover. Validation of the results showed that the model is reasonably effective in predicting both jay abundance and distribution at the extent and resolution used.     

A nutrient balance model (NuBalM) to predict biomass and nitrogen pools in Pinus radiata forests

Forest managers are increasingly required to enhance the productivity and profitability of plantation management while simultaneously reducing the negative ecological effects associated with forest operations. NuBalM (from Nutrient Balance Model) is presented here as a decision support tool that has the potential to assist forest managers in meeting these requirements in Pinus radiata D. Don (radiata pine) plantations. NuBalM incorporates nutrient dynamics and allocation into projections of growth, allowing management techniques to be optimised for productivity and nutrient pool retention over single or multiple rotations.NuBalM was developed using data from biomass, nutrient allocation and soil nutrient dynamics studies conducted in New Zealand radiata pine plantations. The capability of NuBalM to predict stem wood mass based on nitrogen supply and demand was tested against data from multiple trial sites established to examine the effects of variations in stocking, thinning and fertilization regimes. NuBalM satisfactorily predicted stem wood masses across a range of stand ages, with the exception of a trial examining ultra-high applications of nitrogen fertilizer. With the exclusion of the data from this trial, the predicted stem wood masses underestimated the observed figures by a mean value of 1.1 ± 1.0 t ha⁻¹ (95% CI, n = 92).The utility of NuBalM as a tool to predict biomass allocation in radiata pine and nitrogen pools in the forest floor and soil was assessed using comprehensive biomass, nutritional and site data collected from two radiata pine trial sites subjected to differences in organic matter removal at site establishment. NuBalM performed acceptably, generating accurate estimates of stem mass (mean overestimate of 5.5 ± 7.4 t ha⁻¹, 95% CI, n = 6) and total above ground biomass (mean overestimate of 3.1 ± 9.6 t ha⁻¹, 95% CI, n = 6). The effects of organic matter removal and fertilization on total nitrogen pools were also predicted with a reasonable degree of accuracy (mean overestimate of 52 ± 53 kg N ha⁻¹, 95% CI, n = 9).From these results we conclude that NuBalM can be utilised to provide projections of productivity and nitrogen pools in radiata pine plantations, and enables the effects of various management practices to be predicted with a reasonable degree of confidence.     

Detecting interspecific and geographic differentiation patterns in two interfertile oak species (Quercus petraea (Matt.) Liebl. and Q. robur L.) using small sets of microsatellite markers

Genetic analysis was carried out in order to provide insights into differentiation among populations of two interfertile oak species, Quercus petraea and Quercus robur. Gene flow between the two species, local adaptation and speciation processes in general, may leave differential molecular signatures across the genome. Three interspecific pairs of natural populations from three ecologically different regions, one in central Europe (SW Germany) and two in the Balkan Peninsula (Greece and Bulgaria) were sampled. Grouping of highly informative SSR loci was made according to the component of variation they express—interspecific or provenance specific. ‘Species’ and ‘provenance discriminant’ loci were characterized based on F_STs. Locus specific F_STs were tested for deviation from the neutral expectation both within and between species. Data were then treated separately in a Bayesian analysis of genetic structure. By using three ‘species discriminant’ loci, high membership probability to inferred species groups was achieved. On the other hand, analysis of genetic structure based on five ‘provenance discriminant’ loci was correlated with geographic region and revealed shared genetic variation between neighbouring Q. petraea and Q. robur. Small sets of highly variable nuclear SSRs were sufficient to discriminate, either between species or between provenances. Thus, an effective tool is provided for molecular identification of both species and provenances. Furthermore, data suggest that a combination of gene flow and natural selection forms these diversity patterns. ‘Species discriminant’ loci might represent genome regions affected by directional selection, which maintains species identity. ‘Provenance specific’ loci might represent genome regions with high interspecific gene flow and common adaptive patterns to local environmental factors.     

Impact of mistletoes (Amyema miquelii) on host (Eucalyptus blakelyi and Eucalyptus melliodora) survival and growth in temperate Australia

The damaging effects of mistletoes Amyema spp. on host eucalypts in rural Australia concern many landholders, but few data are available to evaluate mistletoe impacts or formulate management strategies. We used an experimental disinfection approach to determine the effects of a putative pest mistletoe, Amyema miquelii, on its two principal hosts, Blakely's red gum (Eucalyptus blakelyi) and yellow box (Eucalyptus melliodora), on pastoral properties in northern New South Wales. For each host species, pairs of nearby trees were matched for heigth, diameter at breast height, level of mistletoe infestation and distance to neighbouring trees. One tree per pair was chosen at random and all the mistletoes pruned. After 33 months all treated trees were still alive, but among control trees seven of 29 red gums and one of 20 yellow box trees had died. The difference in survival between treated and control trees was significant for red gum. Among surviving pairs of trees, treated trees of both species had significantly greater diameter increments and significantly more foliage than untreated trees. After 33 months, the average increase in relative host foliage biomass attributable to removal of mistletoes was 22% in red gum and 24% in yellow box, compared with untreated controls. The average increase in radial growth attributable to mistletoe removal was 55% in red gum and 49% in yellow box. Diameter increment was negatively and linearly related to mistletoe infestation level in control trees of both species. The differences between infected and disinfected trees in this experiment may underestimate the impact of Amyema miquelii on host growth because small amounts of host biomass were unavoidably lost during disinfection. In a separate experiment, loss of host biomass had a significant negative impact on diameter increment. Although the results indicate that control of serious Amyema miquelii infestation of individual trees will be worthwhile in terms of immediate host survival and higher growth rate, prudent long-term management may well allow for the loss of small numbers of farm trees to mistletoes when planning tree recruitment and utilisation.     

Effect of drought on diameter increment of Quercus ilex, Phillyrea latifolia, and Arbutus unedo in a holm oak forest of NE Spain

The present study was carried out to elucidate the drought growth responses of Quercus ilex L., Phillyrea latifolia L., Arbutus unedo L., and other accompanying woody species of the Mediterranean holm oak forest. We submitted holm oak forest stands in Prades mountains (NE Spain) to a 2-year experimental drought. We reduced soil water availability about 15% by plastic strips and funnels that partially excluded rain throughfall and by ditch interception of water runoff. Mean stem diameter increment showed a great variation depending on the species. A. unedo had larger growth rates than Q. ilex and P. latifolia, but it was also the species that experimented the highest growth reduction in the drought plots (77%), suggesting a higher drought sensitivity than Q. ilex (55%) and P. latifolia (no drought effect). The growth reduction was specially marked in the larger trees. Aboveground stand biomass increment, estimated from stem diameter by allometric relationships, was 1.9 Mg ha⁻¹ per year in the control plots. The 15% reduction in the upper soil moisture produced 42% reduction in this biomass increment. In the drier conditions predicted in this Mediterranean area in the frame of climate change, an important reduction of growth rates can be hence expected, accompanied by a gain of dominance of drought-tolerant species such as P. latifolia in detriment of more mesic species such as Q. ilex.     

Effects of treeshelters and polyethylene mulch sheets on survival and growth of cork oak (Quercus suber L.) seedlings planted in northwestern Tunisia

Success of plantation establishment depends on the ability of seedlings to withstand animal browsing and competition from vegetation. This study in northwestern Tunisia examined the influence of browsing protection and vegetation control on seedling mortality, diameter, and height growth components of an evergreen sclerophyllous oak species, cork oak (Quercus suber L.) for the first 2 years after planting (2005 and 2006). Browsing protection was controlled by varying both treeshelter type (non-vented, vented, and control) and height (1.2-m and 1.8-m tall). Competing vegetation was controlled through increasing sizes of square-shaped black polyethylene mulch sheets: 0 m, 0.7 m, 1 m and 1.3 m large. Seedling mortality rate was relatively low during the first year (9%) and was not affected by any of the studied factors. During the following year, mortality was lower for seedlings in both types (non-vented and vented) of 1.2-m tall treeshelters than for unsheltered ones. Mortality was only slightly higher for 1.8-m tall shelters than for 1.2-m shelters. Compared to unsheltered seedlings, basal diameter was reduced during both seasons for seedlings inside non-vented treeshelters (1.2-m and 1.8-m tall), whereas for vented shelters it was reduced during the first year and then increased during the second year. Height growth was positively affected by tree shelters during both years. Seedlings inside non-vented 1.8-m tall shelters were the tallest having an average height of up to 140 cm at the end of the second year, while the unsheltered seedlings had only an average height of up to 42 cm. The greater height of the sheltered seedlings was attributed to the increase of: (1) the number of shoot flushes yearly established on the main stem (up to three), and (2) the length of all the growth units produced during each flush. Seedlings inside vented shelters had balanced growth with a low proportion of seedlings unable to support themselves without a stake (7%). However, non-vented shelters had unbalanced height and diameter growths resulting in both a higher height-to-diameter ratio and in a higher proportion of seedlings having stability problems (47%). Polyethylene mulch sheets did not affect early survival, seedling basal diameter, or number of shoot flushes established yearly, but did slightly improve seedling height at the end of the second season, mostly due to the significant improvement in shoot elongation of the first flush. Results suggest that both 1.8 m treeshelters and plastic mulches may enhance the growth of cork oak seedlings planted on harsh sites in northwestern Tunisia.     

Effects of nutrient addition, mulching and planting-hole size on early performance of Dryobalanops aromatica and Shorea parvifolia planted in secondary forest in Sarawak, Malaysia

Native species of the Dipterocarpaceae are being planted throughout Southeast Asia as a source of future timber and to restore degraded lands. A detailed understanding of factors controlling seedling performance is required for the successful planting of dipterocarps. Below-ground resource availability is hypothesized to have a significant affect on seedling performance of dipterocarp species when planted in selectively logged forests or in open, degraded areas. This study tested three methods thought to increase below-ground resource availability and thereby improve the performance of two dipterocarp species (Dryobalanops aromatica and Shorea parvifolia) when grown in degraded secondary forest: nutrient addition, mulching, and increased planting-hole size. Seedlings of the two species were grown in two planting-hole sizes (12 cm×18 cm and 20 cm×30 cm), with and without nutrients (NPK), and with and without mulching. The experiment was conducted in two sites (Sampadi and Balai Ringin) in Sarawak, Malaysia, to test for spatial variation in treatment effects. Seedling growth and survival were monitored over 22 months. Seedling survival was >94% for both species and did not differ significantly among any of the treatments. High monthly rainfall throughout the first year of the experiment may have enhanced seedling survival. Nutrient addition had the strongest effect on seedling growth for both species, with nutrient addition increasing growth by >50% at Sampadi and for D. aromatica in one block at Balai Ringin. There was significant spatial variation in treatment effects. S. parvifolia did not respond to nutrient addition at Balai Ringin, suggesting that the plants may not have received the added nutrients in that site. Mulching had a positive effect on growth at Sampadi, but no effect or a negative effect at Balai Ringin. Further analysis of the specific effects of mulching on resource supply is required. Planting-hole size did not have a consistent significant effect on the growth of either species. This may have been due to the favorable soil water status during the experiment. The results of this experiment emphasize the potential importance of site effects and interactions between site and treatment effects in enrichment planting trials using dipterocarps. Further experimental studies on below-ground resource limitations of dipterocarp growth in a wide range of sites are required. Studies that monitor the availability and fluxes of soil resources will be the most informative.     

Seedling quality and field performance of commercial stocklots of containerized holm oak (Quercus ilex) in Mediterranean Spain: an approach for establishing a quality standard

Holm oak is the hardwood most used for reforestation in Mediterranean Spain, which makes the development of stock quality standards in order to improve establishment success, a priority. However, its nursery culture is characterized by a wide range of practices resulting in stock heterogeneity and a potentially varied outplanting performance. Previous research has focused on specific seedling quality attributes, obviating the integral effect of nursery culture on overall quality. We studied growing regime, seedling quality, and field performance in nine holm oak stocklots produced in commercial nurseries during two consecutive growing years. Results proved variations in field performance were related to stocklot quality and, hence, to the growing regime practised. This dependence on stock quality may vary with planting site weather: in the drier year, survival was related to attributes like height, water status and K concentration, while, in the second, milder year, only growth performance was related to nutrient concentrations, plant size and water status. Results indicated the following quality standards for height: 12–17 cm, diameter: 3.5–4.8 mm, shoot and root weights: 1.3–1.6 and 2.8–4.7 g, respectively, N–P–K foliar concentrations: over 10–0.9–3.7 mg g⁻¹, respectively and in water status parameters: E_MX < 5 MPa and SWD_TL > 15%. These attributes can be adjusted using nursery cultural practices in order to meet seedling quality standards for holm oak for planting across similar sites.     

Effects of the interaction between drought and shade on water relations, gas exchange and morphological traits in cork oak (Quercus suber L.) seedlings

The combined effect of drought and light on different physiological and biochemical traits was assessed in cork oak (Quercus suber L.) seedlings grown under two levels of light availability and submitted to a long-standing drought. Watering was withdrawn after germination and seedlings were allowed to dry to a water content of ca. 50% of field capacity. At this point, water-stressed seedlings were grown under moderate drought and two light regimes: high light (HL—50%) and low light (LL—2%). Soil water in control plants was kept close to field capacity (90–100%) for both light environments. Water-relations parameters derived from P–V curves, gas exchange and water status at predawn (Ψ_pd) were evaluated at twice during the experiment. Nitrogen and chlorophyll contents were determined in the same leaves used for the gas exchange measurements. In addition, maximum rate of carboxylation (V_cmax) and electronic transport (J_max) were derived from A–C_i curves in well-watered seedlings.

The variation on moisture availability during the experiment was the same under both light environments. In control plants, Ψ_pd was over −0.3 MPa at the two harvests, while stressed seedlings decreased to −0.9 MPa, with no differences between light treatments. Water stress decreased osmotic potentials at full (Ψπ₁₀₀) and zero turgor (Ψπ₀). The regressions between both potentials and Ψ_pd showed a higher intercept in shade grown seedlings. This fact will point out the higher osmoregulation capacity in sun seedlings whatever water availability.

Nitrogen investment on a per leaf mass (N_mass), chlorophyll content (Chl_mass) and SLA tended to show a typical pattern of sun-shade acclimation. Thus, the three parameters increased with shade. Only for N_mass there was a significant effect of watering, since water stress increased N_mass.

LL plants showed a lower photosynthetic capacity in terms of maximum net photosynthesis at saturating light (A_max), which was related to a decrease in V_cmax and J_max. Both parameters varied with specific leaf area (SLA) in a similar way. The low-light environment brought about a higher nitrogen investment in chlorophyll, while under high-light environment the investment was higher in carboxylation (V_cmax) and electronic transport (F_max).

Stomatal conductance to water vapour (g_wv) and A_max were lower in low-light seedlings independently of watering. In addition, there was a trend to keep higher intrinsic water use efficiency (IWUE) under high light environment. The increase of IWUE under water stress was higher in HL seedlings. This was as consequence of the steeper decline in g_wv as Ψ_pd decreased. The decrease of A_max with Ψ_pd occurred in a similar way in LL and HL seedlings. Thus, the HL seedlings tended to sustain a higher ability to increase IWUE than LL seedlings when they were submitted to the same water stress.     

Effects of contour hedgerows on water and soil conservation,crop productivity and nutrient budget for slope farmland in the Three Gorges Region (TGR) of China

Soil erosion has long been recognized as a major environmental problem in the Three Gorges Region (TGR) where slope farming is commonly practiced but the local topography is hilly. In consideration of the poor socioeconomic position of local farmers, low cost hedgerows had been introduced as a soil conservation measure to the TGR in the late 1980s. A collaborative research programme was initiated by the Chinese University of Hong Kong, the Chinese Academy of Science, the Huazhong Agricultural University, and the Bureau of Soil and Water Conservation of Zigui County to study the potential of adopting hedgerows in the TGR. Six experimental plots (10 m × 2 m, gradient = 25°) were constructed at Zigui County, Hubei Province to study effects of hedgerows on erosion, nutrient loss and crop productivity. Results indicated that there were significant relationships between rainfall and runoff, and rainfall and soil loss, respectively. Conventional slope farming could not be considered a sustainable agricultural practice because it resulted in severe erosion and low crop yield. Hedgerows per se seemed not to be effective in reducing soil loss and boosting crop productivity, but performances could be greatly improved when they combined with the use of fertilizers. Current farming and fertilization practices, however, generally did not meet N demands of crops. Results and findings of this paper will contribute towards a technical reference for the promotion and adoption of hedgerows in the TGR.     

Effect of spacing on growth and quality parameters in sessile oak (Quercus petraea) stands in central Poland: results 7 years after planting

We examined the growth and quality of young oaks in stands established at three different spacings. The three densities were the traditional level, that is, 8000?seedlings?ha^?1 (1.5?×?0.85?m) plus two where the initial density was decreased by 50% and 70%, achieved by increasing the distance between the rows from 1.5 to 3 or 4.5?m. The experiment was located in central Poland on a moderately fertile site. In the two wider spacings, the intention was to use natural regeneration occurring between oak rows as a replacement of intraspecific competition for oaks. The mean height was only affected in the third and the fourth year after establishment. Subsequently, the differences between spacings with respect to both growth and quality traits were insignificant and small. At lower densities, natural regeneration of willow and aspen was abundant and inter-species competition replaced intra-species competition, thus explaining the absence of a spacing effect on growth and quality properties. In the study period, the wider spacings were economically beneficial compared to traditional densities. A strong competition affecting height growth caused by natural regeneration necessitated extensive tendings. More tendings are likely to be needed in the future and they might reduce the initial economical superiority of wider spacing. The study indicates that it is possible to decrease the initial spacing in oak plantations and simultaneously support good growing conditions by means of natural processes, for example, natural regeneration. However, economic efficiency of the method has to be studied in a longer perspective.     

Nutrient store and export rates of Eucalyptus urograndis plantations in eastern Amazonia (Jari)

Nutrient loss due to export of wood and bark of 4.5-year-old Eucalyptus urograndis was studied in the Jari project, eastern Amazonia, Brazil. Results are based on estimates of biomass, above-ground nutrient store, and nutrient stores in the soil. Thirteen stands varying in site conditions and the number of previous rotations (up to three), were studied. Average nutrient losses due to export of wood and bark, as a percentage of the element store in the above-ground tree biomass, were: N, 65%, P, 54%; Ca, 76%; K, 57%; Mg, 61%. A comparison of export losses with the present soil stores of Ca, K and Mg cations shows that export is equal to or surpasses the remaining soil store to a depth of 100 cm. As a result of the last narvest, about 200–250 kg Ca and about 100 kg K ha⁻¹ were removed. In spite of uncertainties about other unknown inputs and outputs, one has to conclude that the next rotation may be endangered by acute deficiency of mineral nutrients, especially of Ca.     

Susceptibility of Eucalyptus globulus ssp. globulus to sawfly (Perga affinis ssp. insularis) attack and its potential impact on plantation productivity

Sawflies (Perga species) are leaf-damaging pests of the major plantation tree species Eucalyptus globulus ssp. globulus. This work describes the patterns of quantitative genetic variation in susceptibility to attack by Perga affinis ssp. insularis, based on data from a large open-pollinated trial containing genetic material from throughout the geographic range of E. globulus ssp. globulus. Forty three per cent of the trees in the trial exhibited damage from sawflies in either their sixth or seventh growth seasons. The incidence of damage was genetically based, with significant variation between geographic races of E. globulus ssp. globulus and highly heritable (h²=0.43±0.05) variation in damage incidence within races. Susceptibility to sawfly damage significantly affects plant fitness with both genetic and environmental correlations demonstrating that sawfly damage causes slow growth and increased mortality of trees. Mild and severe sawfly damage resulted in 16 and 31% reduction in the basal area of surviving trees, and the effect was consistent across races and families. Based on this data, we propose a model predicting relative plantation productivity of races of different susceptibility under different infestation regimes.     

Effects of tending practices on nutrient dynamics in a young stand of sugi (Cryptomeria japonica) and hinoki (Chamaecyparis obtusa)

The effects of three silviculture practices, fertilization, weeding, and pruning, on nutrient losses and cycling were investigated for 8 years in a young coniferous stand and compared with an established stand. On the basis of the results, the effects of three intensive operations on site productivity are discussed. The quantities of organic matter and nutrients supplied by weeding and pruning were large, these two operations having a beneficial effect on nutrient cycling and site productivity before crown closure in the young stand. Evapotranspiration was estimated to be high in the young stand before crown closure and water loss (precipitation - discharge) decreased after pruning to approximately 25% of the prepruning value. The effects of fertilization on stream-water quality were restricted to the period directly after clearcutting. There was a positive relationship between stream nutrient export and stream discharge except for the first 3 years after clearcutting and planting. To maintain site productivity it is important to prevent excess nutrient loss in stream discharge.     

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.