Can we use shelterwoods in Mediterranean pine forests to promote oak seedling development?

The use of shelterwoods to favour the development of natural or underplanted seedlings is common in temperate forests but rare in the pine forests of the Mediterranean area. Our aim was to assess the use of shelterwoods in Aleppo pine (Pinus halepensis) woodlands in southern France to promote the survival and growth of two co-occurring oak species: the deciduous Quercus pubescens and the evergreen Quercus ilex.Twelve Aleppo pine stands were selected and differentially thinned to create a light shelterwood (basal area = 10 m²/ha, irradiance 52%), a medium shelterwood (basal area = 19 m²/ha, irradiance 33%) and a dense shelterwood (basal area = 32 m²/ha, irradiance 13%). A total of 1248 sowing points, half composed of Q. pubescens and half of Q. ilex, were then set up in these three conditions. Seedling survival and growth were monitored for 3 years. Plant stress was assessed by measuring predawn leaf potential and photosynthetic performance through the F_v/F_m ratio. Soil moisture was also recorded at two depths during two growing seasons.Survival was high for both species in all three conditions due to three consecutive wet years. The lowest survival was recorded for Q. pubescens in the dense shelterwoods. Growth in diameter and height increased from the dense to the light shelterwoods. Shrubs developed more strongly in the light shelterwood, and increasing shrub cover enhanced height growth. Photosynthetic performance was lowest for Q. pubescens in dense shelterwoods and highest in light shelterwoods, whereas the reverse was true for Q. ilex. The lowest predawn potentials were recorded in the dense shelterwoods even though higher soil water content values were measured in this treatment during the summer drought.We show that light shelterwoods were more beneficial to growth than denser ones, indicating control mainly by light availability during the 3 years of the study. However, as lower soil moisture at 30-50 cm depth and faster understorey development were also recorded in this condition, more extended observation is needed to determine whether this benefit persists in subsequent years.     

Water use by Tabor and Kermes oaks growing in their respective habitats in the Lower Galilee region of Israel

We estimated water use by the two main oak species of the Lower Galilee region of Israel—Tabor (Quercus ithaburensis) and Kermes (Quercus calliprinos)—to develop management options for climate-change scenarios. The trees were studied in their typical phytosociological associations on different bedrock formations at two sites with the same climatic conditions. Using the heat-pulse method, sap flow velocity was measured in eight trunks (trees) of each species during a number of periods in 2001, 2002 and 2003. Hourly sap flux was integrated to daily transpiration per tree and up-scaled to transpiration at the forest canopy level. The annual courses of daytime transpiration rate were estimated using fitted functions, and annual totals were calculated. Sap flow velocity was higher in Tabor than in Kermes oak, and it was highest in the youngest xylem, declining with depth into the older xylem. Average daytime transpiration rate was 67.9 ± 4.9 l tree⁻¹ d⁻¹, or 0.95 ± 0.07 mm d⁻¹, for Tabor oak, and 22.0 ± 1.7 l tree⁻¹d⁻¹, or 0.73 ± 0.05 mm d⁻¹, for Kermes oak. Differences between the two oak species in their forest canopy transpiration rates occurred mainly between the end of April and the beginning of October. Annual daytime transpiration was estimated to be 244 mm year⁻¹ for Tabor oak and 213 mm year⁻¹ for Kermes oak. Adding nocturnal water fluxes, estimated to be 20% of the daytime transpiration, resulted in total annual transpiration of 293 and 256 mm year⁻¹ by Tabor and Kermes oaks, respectively. These amounts constituted 51% and 44%, respectively, of the 578 mm year⁻¹ average annual rainfall in the region. The two species differed in their root morphology. Tabor oak roots did not penetrate the bedrock but were concentrated along the soil–rock interface within soil pockets. In contrast, the root system of Kermes oak grew deeper via fissures and crevices in the bedrock system and achieved direct contact with the deeper bedrock layers. Despite differences between the two sites in soil–bedrock lithological properties, and differences in the woody structure, annual water use by the two forest types was fairly similar. Because stocking density of the Tabor oak forests is strongly related to bedrock characteristics, thinning as a management tool will not change partitioning of the rainfall between different soil pockets, and hence soil water availability to the trees. In contrast, thinning of Kermes oak forests is expected to raise water availability to the remaining trees.     

Canopy gaps and regeneration in old-growth Oriental beech (Fagus orientalis Lipsky) stands, northern Iran

Virgin beech Fagus orientalis forests in northern Iran provide a unique opportunity to study the disturbance regimes of forest ecosystems without human influence. The aim of this research was to describe characteristics of natural canopy gaps and gap area fraction as an environmental influence on the success of beech seedling establishment in mature beech stands. All canopy gaps and related forest parameters were measured within three 25 ha areas within the Gorazbon compartment of the University of Tehran’s Kheyrud Experimental Forest. An average of 3 gaps/ha occurred in the forest and gap sizes ranged from 19 to 1250 m² in size. The most frequent (58%) canopy gaps were <200 m². In total, canopy gaps covered 9.3% of the forest area. Gaps <400 m² in size were irregular in shape, but larger gaps did not differ significantly in shape from a circle. Most gaps (41%) were formed by a single tree-fall event and beech made up 63% of gap makers and 93% of gap fillers. Frequency and diversity of tree seedlings were not significantly correlated with gap size. The minimum gap size that contained at least one beech gap-filling sapling (<1.3 m tall) was 23.7 m². The median gap size containing at least one beech gap-filling sapling was 206 m² and the maximum size was 1808 m². The management implications from our study suggest that the creation of small and medium sized gaps in mixed beech forest should mimic natural disturbance regimes and provide suitable conditions for successful beech regeneration.     

Growth and nutrition of Pinus radiata in response to fertilizer applied after thinning and interaction with defoliation associated with Essigella californica

Infestations of Essigella californica following the installation of post-thinning fertilizer trials in Pinus radiata plantations provided an opportunity to examine the impact of repeated defoliation over a period of 8 years (1997–2005). Replicated treatments (n = 4) of nil fertilizer (control), N (300 kg ha⁻¹) as urea, P (80 kg ha⁻¹) and S (45 kg ha⁻¹) as superphosphates were applied immediately after thinning at three sites and this was followed by a second application of NPS fertilizers 6 years later with N applied at 300 kg ha⁻¹ as urea and ammonium sulphate and P at 80 or 120 kg ha⁻¹. Defoliation of untreated P. radiata gradually increased to 50% over a period of 8 years. Basal area growth was negatively correlated with average defoliation for two consecutive post-fertilizer periods of 6 and 2 years. Growth responses to fertilizer varied considerably between sites but the largest improvement in growth was due to NPS fertilizer, this increased basal area by 30–80%. Application of N fertilizer raised total N levels in foliage and increased defoliation with a commensurate loss in growth under conditions of deficiencies of S or P. Repeated infestations gradually increased the percentage of trees with severe defoliation (>80% loss of foliage) indicating that nutrient-deficient trees have a reduced capacity for foliage recovery between episodes of peak infestation. In contrast, treatment with N fertilizer in combination with S- and P-corrected deficiencies of these nutrients, raised levels of total N in foliage and reduced defoliation to approximately 20%. Basal area growth responses to NPS fertilizers reflected improved nutrition as well as reduced insect damage. The reduction in defoliation under conditions of balanced tree nutrition was most likely due to enhanced needle retention following correction of P deficiency as well as greater availability of nutrients enabling a more vigorous recovery of P. radiata after an episode of E. californica activity. Treatment with fertilizer therefore reduced the long-term impact of aphid damage and improved growth of P. radiata.     

Enrichment planting as a silvicultural option in the eastern Amazon: Case study of Fazenda Cauaxi

Liana-dominated forest patches constitute 15–20% of old-growth forests in the Eastern Amazon but are generally excluded from management for timber production. Here we ask if liana-dominated patches may be brought into production by clearing lianas and conducting enrichment planting (EP) of native timber species. We present growth results from 8 years of such EP trials. Rapid growth and low mortality of all species in this study suggest that EP in cleared liana patches can contribute to timber stocks in second and third harvests of managed forests. The most vigorous individuals of Parkiagigantocarpa and Schizolobium amazonicum in each enrichment site grew more than 1 cm diameter per year (rates were initially >2 cm yr⁻¹), and attained dominant canopy positions and diameters equal to those of small canopy trees in the surrounding forest within 8 years of planting (mean dbh ∼18 cm and ∼20 cm, respectively, at year 8). Limited data on Ceiba pentandra plantings indicate a similar trajectory for this species (dbh ∼40 cm in 8 years). The most vigorous Swietenia macrophylla grew at least 1 cm per year in enrichment plots (mean dbh ∼10 cm in 8 years), but take longer to attain dominant positions. Tabebuia serratifolia may take much longer to reach the canopy than other species tested (rates <1 m yr⁻¹). We attribute the excellent performance to light availability; planting in intact soil with minimal compaction and abundant organic material; and low competition rates maintained by periodic thinning of competing vegetation.     

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.     

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.     

Optimizing the joint production of timber and bilberries

Berries and mushrooms are increasingly appreciated products of Finnish forests. Therefore, there is a need to integrate them in silvicultural planning. Bilberry (Vaccinium myrtillus L.) is an economically important wild berry that is widely collected for household consumption and sale in North Karelia, Finland. In this study, bilberry yield models developed recently were included in a stand growth simulator and the joint production of timber and bilberry was optimized by maximizing soil expectation value (SEV) with 3% discounting rate, assuming that 75% of the bilberry yield is harvested. The effect of bilberry production on the optimal stand management increased with increasing bilberry price. With high bilberry prices (4–8 € kg⁻¹) it was optimal to manage the mixed stand of Scots pine, Norway spruce and birch, and the pure stand of Norway spruce so as to promote bilberry production. In the Scots pine stand, where bilberry yields are higher, bilberry production affected optimal stand management already with a price of 2 € kg⁻¹. Compared to timber production, joint production led to longer rotation lengths, higher thinning intensities, more frequent thinnings, and higher share of Scots pine in the mixed stand. The contribution of bilberries to the total SEV increased with increasing bilberry price and discounting rate. In the mixed stand and pine stand the SEV of bilberry production, calculated with 3% discounting rate, exceeded the SEV of timber production when bilberry price was 4 € kg⁻¹.With 4% discounting rate this happened already with bilberry price of 2 € kg⁻¹. It was concluded that forest management which promotes bilberry yields is the most profitable in pine stands where the potential bilberry yields are high.     

Compositional changes of forest-floor vegetation in young stands of Norway spruce as an effect of repeated fertilisation

Forestry practices that aim to increase biomass production may mitigate climate change through increased carbon sequestration and the potential of substituting fossil fuels with renewable biofuels. Fertilising young stands of Norway spruce in Sweden have shown to increase tree growth by more than 200%. Fertilisation, however, also has other effects on forest ecosystems. Here, we studied the response of the species composition of forest-floor vegetation to three different frequencies of fertilisation in young stands of Norway spruce. Fertiliser was applied every year, every second year or every third year. The total amount of N ranged from 425 kg ha⁻¹ to 625 kg ha⁻¹, in combination with P, K, Ca, Mg, S, Mn, Zi, B and Cu. The largest effects of the fertilisation were found among bryophytes and lichens, which lost substantial cover. Unexpectedly, Deschampsia flexuosa, commonly known to be favoured by fertilisation, was negatively affected. Species that increased in frequency were Oxalis acetosella, Brachythecium sp. and Plagiothecium sp. Decreased availability of light, as an indirect effect of fertilisation through increased tree canopy cover, was found to be the most important factor behind the change in species composition of vascular plants. The total cover of bryophytes, however, did not show any significant response to the changes in canopy cover, indicating that the effects seen in this group may be a result of more direct effects of the fertiliser. Few significant differences were found between the two most intensive fertilisation frequencies, although fertilisation every third year was often distinguished from both the control and the other fertilised treatments. Even though the effects at the stand level were substantial, the effects on biodiversity and function of ecosystems on a landscape or regional level need further investigation.     

Effects of [CO2] and nitrogen on morphological and biomass traits of white birch (Betula papyrifera) seedlings

To investigate the interactive effects of CO₂ concentration ([CO₂]) and nitrogen supply on the growth and biomass of boreal trees, white birch seedlings (Betula papyrifera) were grown under ambient (360 μmol mol⁻¹) and elevated [CO₂] (720 μmol mol⁻¹) with five nitrogen supply regimes (10, 80, 150, 220, and 290 μmol mol⁻¹) in greenhouses. After 90 days of treatment, seedling height, root-collar diameter, biomass of different organs, leaf N concentration, and specific leaf area (SLA) were measured. Significant interactive effects of [CO₂] and N supply were found on height, root-collar diameter, leaf biomass, stem biomass and total biomass, stem mass ratio (SMR), and root mass ratio (RMR), but not on root mass, leaf mass ratio (LMR), leaf to root ratio (LRR), or leaf N concentration. The CO₂ elevation generally increased all the growth and biomass parameters and the increases were generally greater at higher levels of N supply or higher leaf N concentration. However, the CO₂ elevation significantly reduced SLA (13.4%) and mass-based leaf N concentration but did not affect area-based leaf N concentration. Increases in N supply generally increased the growth and biomass parameters, but the relationships were generally curvilinear. Based on a second order polynomial model, the optimal leaf N concentration was 1.33 g m⁻² for height growth under ambient [CO₂] and 1.52 g m⁻² under doubled [CO₂]; 1.48 g m⁻² for diameter under ambient [CO₂] and 1.64 g m⁻² under doubled [CO₂]; 1.29 g m⁻² for stem biomass under ambient [CO₂] and 1.43 g m⁻² under doubled [CO₂]. The general trend is that the optimal leaf N was higher at doubled than ambient [CO₂]. However, [CO₂] did not affect the optimal leaf N for leaf and total biomass. The CO₂ elevation significantly increased RMR and SMR but decreased LMR and LRR. LMR increased and RMR decreased with the increasing N supply. SMR increased with increase N supply up to 80 μmol mol⁻¹ and then leveled off (under elevated [CO₂]) or stated to decline (under ambient [CO₂]) with further increases in N supply. The results suggest that the CO₂ elevation increased biomass accumulation, particularly stem biomass and at higher N supply. The results also suggest that while modest N fertilization will increase seedling growth and biomass accumulation, excessive application of N may not stimulate further growth or even result in growth decline.     

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

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

Variation in leaf nitrogen and phosphorus stoichiometry in Picea abies across Europe: An analysis based on local observations

Nitrogen (N) and phosphorus (P) and N:P ratio in terrestrial plants and the patterns at a large geographical scale are an important issue in ecological stoichiometry. In particular, it is essential to know that for a single species, how the N:P stoichiometry varies with climatic factors in the context of global warming. Our analysis was based on a data set including 2583 observations at 441 sites on nutritional status of Norway spruce (Picea abies L.) located in European counties (including Austria, Belgium, Bulgaria, Czech Rep., Finland, Germany, Ireland, Italy, Lithuania, Norway, Slovak Rep., Slovenia, United Kingdom). Our objectives are to demonstrate how leaf N and P concentration and N:P ratio in Norway spruce vary with altitude (ALT), latitude (LAT), longitude (LON), mean annual temperature (MAT) and mean annual precipitation (MAP) across Europe. The results showed that for 1-year-old needles of Norway spruce, the N and P concentration were 13.28 mg g⁻¹, 1.41 mg g⁻¹ and the N:P ratio was 9.76. Leaf N displayed a convex curve pattern with increasing MAT and decreasing LAT from the boreal Europe to the Mediterranean area. The N concentration and N:P generally reached peak at about 7 °C in MAT or 53° N in LAT. The N:P ratio varied non-linearly with LAT and MAP, but linearly with MAT. Leaf N concentration and N:P ratio decreased linearly with increasing ALT in temperate European area. Across Europe, that the patterns of leaf N and N:P ratio were mainly driven by climate-related geochemistry and plant physiology, but also greatly impacted by anthropogenic N deposition.     

A positive growth response to NaCl applications in Eucalyptus plantations established on K-deficient soils

Contrasting responses of Eucalyptus trees to K fertilizer applications have been reported on soils with low K contents. A complete randomized block experiment was set up in Brazil to test the hypothesis that large atmospheric deposits of NaCl in coastal regions might lead to a partial substitution of K by Na in Eucalyptus physiology and enhance tree growth. Treatments with application of 1.5, 3.0, 4.5 kmol K ha⁻¹ (K_1.5, K_3.0, K_4.5, respectively) as KCl, 3.0 kmol K ha⁻¹ applied as K₂SO₄, 3.0 kmol Na ha⁻¹ (Na_3.0) as NaCl commercialized for cattle feeding, and a mixture of 1.5 kmol K + 1.5 kmol Na ha⁻¹ (K_1.5 + Na_1.5) were compared to a control treatment (C) with no K and Na applications. All the plots were fertilized with large amounts of the other nutrients.     

Mixed-species plantations of Acacia mangium and Eucalyptus grandis in Brazil: 1. Growth dynamics and aboveground net primary production

Nitrogen fertilizer inputs increased sharply over the last decade in Brazilian eucalypt plantations. Due to the economic and potential environmental cost of fertilizers, mixed plantations with N-fixing species might be an attractive option to improve the long-term soil N status. A randomized block design was set up in southern Brazil, including a replacement series and an additive series design, as well as a nitrogen fertilization treatment. The development of mono-specific stands of Eucalyptus grandis (0A:100E) and Acacia mangium (100A:0E) was compared with mixed plantations in proportions of 1:1 (50A:50E), and other stands with different densities of acacia for the same density of eucalypts. The objective was to assess the effect of inter-specific interactions on the early development of the two species. Aboveground biomass was measured 6, 12, 18 and 30 months after planting, sampling 6–10 trees of each species per treatment at each age, and allometric equations were established in 0A:100E, 100A:0E, 50A:50E and 50A:100E. The height and basal area of E. grandis seedlings were enhanced by 12% and 30%, respectively by N fertilization at age 1 year. Inter-specific competition led to a stratified canopy, with suppression in acacia growth earlier for basal area than for height. The mean number of stems per acacia tree at 36 months after planting was significantly higher in pure stands (3.7), than in 50A:50E (2.7) and in the additive series (between 1.6 and 1.8). H/D ratios were highly sensitive to inter-tree competition for the two species. The suppressed acacia understorey in mixed-species stands did not influence biomass production and partitioning within eucalypts. This pattern led to biomass accumulation combining the two species in 50A:100E that was about 10% higher than in 0A:100E, from age 12 months onwards. Aboveground net primary production (ANPP) amounted to 25 Mg ha⁻¹ and 37 Mg ha⁻¹ from age 18 to 30 months in 100A:0E and 0A:100E, respectively. Acacia ANPP in 50A:100E amounted to 2 Mg ha⁻¹ over the same period, as a result of substantial inter-specific competition. An increment in biomass production in these very fast-growing eucalypt plantations was achieved introducing acacia as an understorey and not in the 50A:50E design, as observed in other studies.     

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.     

Effect of species and spacing of fast-growing nurse trees on growth of an indigenous tree, Hopea odorata Roxb., in northeast Thailand

We tested the effects of species and spacing of nurse trees on the growth of Hopea odorata, a dipterocarp tree indigenous to Southeast Asia, in a two-storied forest management system in northeast Thailand. Eucalyptus camaldulensis, Acacia auriculiformis, and Senna siamea were planted as nurse trees in 1987 at spacings of 4 m × 8 m, 2 m × 8 m, 4 m × 4 m, and 2 m × 4 m in the Sakaerat Silvicultural Research Station of the Royal Forest Department, Thailand. Seedlings of H. odorata were planted in the nurse tree stands at a uniform spacing of 4 m × 4 m and in control plots (no nurse trees) in 1990. Stem numbers of some nurse trees were thinned by half in 1994. The stem diameter and height of all trees were measured annually until 1995 and again in 2007. The mean annual increment (MAI) in volume was estimated as 8.2–10.1 m³ ha⁻¹ year⁻¹ for E. camaldulensis and 0.9–1.2 m³ ha⁻¹ year⁻¹ for S. siamea, smaller than reported elsewhere. This suggests that the site properties were not suitable for them. The MAI of A. auriculiformis was 7.9–9.8 m³ ha⁻¹ year⁻¹, within the reported range. Survival rates of H. odorata in the S. siamea stands and the control plots decreased rapidly during the first 2 years but then stayed constant from 1992. In contrast, survival rates of H. odorata in the E. camaldulensis and A. auriculiformis stands were initially high (>70%), but then decreased after 1995. Stem diameter, tree height, and stand basal area of H. odorata were large in both the S. siamea stands and the control plots from then. The growth of H. odorata was largest in the 2 m × 8 m S. siamea stands. In contrast, it was restricted in the E. camaldulensis and A. auriculiformis stands owing to strong shading by their canopies. Thinning by 50% tended to facilitate the growth of H. odorata temporarily in the E. camaldulensis and A. auriculiformis stands. The stand basal areas of nurse trees and of H. odorata showed a trade-off. These results suggest that the growth of H. odorata was maximized in the S. siamea stands. We assume, however, that the growth of H. odorata could be improved even in the E. camaldulensis and A. auriculiformis stands by frequent or heavy thinning.     

Patterns of spread of Pinus contorta Dougl. ex Loud. invasion in a Natural Reserve in southern South America

Pinus species, which have formed the foundation of commercial forestry industry in many countries, are known to be invasive in natural ecosystems, especially in the Southern Hemisphere. Pinus contorta is considered one of the most aggressively invasive plantation species. In this paper we aim to: (a) determine patterns of P. contorta invasion in relation to its size and age structure and spatial distribution and (b) determine the effect of vegetation cover on its regeneration. For this purpose, we have chosen the Malalcahuello National Reserve in south-central Chile, which is dominated by Araucaria araucana forests. In the three P. contorta trial plots, attributes were measured in order to describe its current structure. We selected one of the three P. contorta trial plots and set eight 50 m wide and 125 m centrifugal transects starting at the North Azimuth (0°) orientation and then one each 45°. In each transect 25 circular plots of 2.5 m radius were established every 25 m. In each plot, we measured collar diameter (CD) for each P. contorta individual and adjusted a diameter–age function. We recorded the presence of cones for each individual P. contorta within the plots and the total number of P. contorta individuals. In each plot, we measured percent vegetation cover (grass, understory and canopy). The selected parent stand showed a decrease in density from the original plantation spacing, from 2500 to 150 plants ha⁻¹. Of all sampled individuals, only a 38% had cones. We found reproductive structures in trees as young as 5-year-old. Regeneration was found in all transects. Mean density for the area was 1600 plants ha⁻¹, and the greater plant number was found in the West transect, reaching an average of 6600 plants ha⁻¹. Through the interpolation performed with the kriging method, a map of the area with the spatial gradient of plant density was obtained. Naturally regenerated individuals of P. contorta occupied an area of 78 ha where the most distant individual is located in the southeast direction at an average distance of 1200 m from the three study plots. We determined that a positive association exists between P. contorta and the species A. araucana and Nothofagus antarctica. The capacity for early and consistent reproduction and the establishment of individuals dispersed at great distances from the original plots allow us to conclude that P. contorta has great potential as an invasive species in forests of this area of the Andes of southern South America. Invasion of P. contorta has many important implications for the conservation of native forests in our region including diminished regeneration of A. araucana.     

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 changes induced by Acacia mangium plantation establishment: Comparison with secondary forest and Imperata cylindrica grassland soils in South Sumatra,Indonesia

Acacia plantation establishment might cause soil acidification in strongly weathered soils in the wet tropics because the base cations in the soil are translocated rapidly to plant biomass during Acacia growth. We examined whether soils under an Acacia plantation were acidified, as well as the factors causing soil acidification. We compared soils from 10 stands of 8-year-old Acacia mangium plantations with soils from 10 secondary forests and eight Imperata cylindrica grasslands, which were transformed into Acacia plantations. Soil samples were collected every 5–30 cm in depth, and pH and related soil properties were analyzed. Soil pH was significantly lower in Acacia plantations and secondary forests than in Imperata grasslands at every soil depth. The difference was about 1.0 pH unit at 0–5 cm and 0.5 pH unit at 25–30 cm. A significant positive correlation between pH and base saturation at 0–20 cm depth indicated that the low pH under forest vegetation was associated with exchangeable cation status. Using analysis of covariance (ANCOVA), with clay content as the covariate, exchangeable Ca (Ex-Ca) and Mg (Ex-Mg) stocks were significantly lower in forested areas than in Imperata grasslands at any clay content which was strongly related to exchangeable cation stock. The adjusted average Ex-Ca stock calculated by ANCOVA was 249 kg ha⁻¹ in Acacia plantations, 200 kg ha⁻¹ in secondary forests, and 756 kg ha⁻¹ in Imperata grasslands at 0–30 cm. Based on a comparison of estimated nutrient stocks in biomass and soil among the vegetation types, the translocation of base cations from soil to plant biomass might cause a decrease in exchangeable cations and soil acidification in Acacia plantations.     

Forest stand dynamics and sudden oak death: Mortality in mixed-evergreen forests dominated by coast live oak

Sudden oak death (SOD), caused by the recently discovered non-native invasive pathogen, Phytophthora ramorum, has already killed tens of thousands of native coast live oak and tanoak trees in California. Little is known of potential short and long term impacts of this novel plant–pathogen interaction on forest structure and composition. Coast live oak (Quercus agrifolia) and bay laurel (Umbellularia californica) form mixed-evergreen forests along the northern California coast. This study measured tree mortality over a gradient of disease in three time periods. Direct measurements of current mortality were taken during 2004, representing a point-in-time estimate of present and ongoing mortality. Past stand conditions, c. 1994, were estimated using a stand reconstruction technique. Future stand conditions, c. 2014, were calculated by assuming that, given a lack of host resistance, live trees showing signs of the disease in 2004 would die. Results indicate that coast live oaks died at a rate of 4.4–5.5% year⁻¹ between 1994 and 2004 in highly impacted sites, compared with a background rate of 0.49% year⁻¹, a ten-fold increase in mortality. From 2004 to 2014, mortality rates in the same sites were 0.8–2.6% year⁻¹. Over the entire period, in highly impacted sites, a 59–70% loss of coast live oak basal area was predicted, and coast live oak decreased from 60% to 40% of total stand basal area, while bay laurel increased from 22% to 37%. Future stand structures will likely have greater proportions of bay laurel relative to coast live oak.