Long-term trends in loblolly pine productivity and stand characteristics in response to thinning and fertilization in the West Gulf region

Two levels each of thinning and fertilization were applied to a 7-year-old loblolly pine (Pinus taeda L.) plantation on a nitrogen- and phosphorus-deficient West Gulf Coastal Plain site in Louisiana. Levels of thinning were no thinning, or thinning applied 7 and 14 years after stand initiation. Levels of fertilization were no fertilization or broadcast fertilization with diammonium phosphate at age 7 years plus refertilization with urea, monocalcium phosphate, and potash at age 14 years. Long-term measurements of climate, stand development and productivity, projected leaf area index, and foliar nutrition were initiated at age 11 years. We found that by age 17 years, thinning increased mean live-crown length from 4.2 to 7.8 m, and mean tree diameter from 15.0 to 21.8 cm compared to the unthinned treatment. After rethinning at age 14 years, stand basal area increased 1.2 and 19.2% between ages 15 and 17 years on the unthinned and thinned plots, respectively. Refertilization at age 14 years reestablished foliar N, P and K sufficiency, which increased leaf area index from 4.2 to 6.0 m² m⁻² on the unthinned plots and from 3.2 to 3.8 m² m⁻² on the thinned plots, and subsequently, increased gross stand biomass from 114 to 141 Mg ha⁻¹ on the unthinned plots and from 78 to 95 Mg ha⁻¹ on the thinned plots by age 17 years. Leaf area was an important factor controlling loblolly pine productivity. At our study site, however, competition for light and water and nutrition-limited foliage growth influenced the variability and scope of this relationship. Our results suggest that a positive and linear relationship between leaf area and loblolly pine productivity does not universally occur on loblolly pine sites.     

Long-term growth and ecophysiological responses of a southeastern Oklahoma loblolly pine plantation to early rotation thinning

A thinning levels study was initiated in a 9-year-old loblolly pine (Pinus taeda L.) plantation containing 26.6 m² ha⁻¹ basal area during the spring of 1984 in southeastern Oklahoma. Thinning treatments consisted of (1) three control plots (BA100), (2) three plots thinned to approximately 50% of the original basal area (BA50) and (3) three plots that were thinned to 25% of the original basal area (BA25). In 1987 the BA50 and BA25 plots were both rethinned to a basal area of 12 m² ha⁻¹. No other thinnings were done through age 24.The control plots have attained a basal area of 45.3 m² ha⁻¹ and basal area is now starting to decline. The BA25 and BA50 plots have basal areas between 34 and 35 m² ha⁻¹. Mortality has averaged about 90 trees ha⁻¹ per year from age 10 to age 24 on the control plot, declining from 2078 trees ha⁻¹ at age 10 to 827 trees ha⁻¹ at age 24. Mortality losses in the BA25 and BA50 plots have been only 3.2–7.7 trees ha⁻¹ per year over the entire study period. Cumulative stem biomass lost to mortality was 10.5, 16.0 and 61 Mg ha⁻¹, respectively, for the BA25, BA50 and BA100 treatments. Cumulative standing live biomass at age 24 in the BA100 treatment is 132 Mg ha⁻¹. Cumulative standing live biomass in the BA25 and BA50 treatments at age 24 is 86 and 79%, respectively, of that observed in the BA100 treatment. These results suggest wide ranges of residual stand densities left after an early thinning will produce a high percentage of the potential total maximum standing stem biomass. Diameter distributions at age 24 show only 33% of the trees in the BA100 treatments have the dimensions to be sawtimber (≥30 cm) but 92 and 95% of the trees in the BA25 and BA50, respectively, are sawtimber dimension or larger. Mean annual stem biomass production (MAI) of the BA100 treatment is 7.5 Mg ha⁻¹ per year at age 24. MAI of the thinned treatments is about 5.1 Mg ha⁻¹ per year and is converging to that of the BA100 treatment. The basis for this convergence is not that the live trees in the BA100 treatment are producing live biomass less rapidly than the thinned plots, but that mortality losses in the BA100 plot are much higher. Current annual stemwood production in all treatments is often limited by the severe summer droughts that occur in this region. The wide variations in weather experienced at this site also result in variations in earlywood:latewood ratio and ring specific gravity.     

Stand development after different thinnings in two uneven-aged Picea abies forests in Sweden

Two field experiments, located in Central and Northern Sweden, were used to study the influence of standing volume on volume increment and ingrowth in uneven-aged Norway spruce (Picea abies (L.) Karst.) stands subjected to different thinnings. Each experiment had a 3 × 2 factorial block design with two replications. Treatments were thinning grade, removing about 45, 65, and 85% of pre-thinning basal area, and thinning type, removing the larger or the smaller trees, respectively. Each site also had two untreated control plots. Plot size was 0.25 ha. Volume increment was 0.5–6.8 m³ ha⁻¹ year⁻¹ for the plots, and significantly positively (p < 0.01) correlated with standing volume. Within treatment pairs, plots thinned from Above had consistently higher volume increment than plots thinned from Below. Ingrowth ranged from 3 to 33 stems ha⁻¹ year⁻¹, with an average of 14 and 21 stems ha⁻¹ year⁻¹ at the northern and southern site, respectively. At the southern site ingrowth was significantly negatively (p < 0.01) correlated with standing volume, but not at the northern site. Mean annual mortality after thinning was 2 and 7 stems ha⁻¹ year⁻¹at the northern and southern site, respectively.     

Stand development and production dynamics of loblolly pine under a range of cultural treatments in north-central Florida USA

The objectives of this study were to examine the effects of stand development and soil nutrient supply on processes affecting the productivity of loblolly pine (Pinus taeda L.) over a period approximately equal to a pulpwood rotation (18 years). The experiment consisted of a 2×2 factorial combination of complete and sustained weed control and annual fertilization treatments (C: control treatment, F: fertilization, W: weed control, FW: combined fertilization and weed control), located on a Spodosol in north-central Florida, USA. The reduction of soil nutrient limitations through fertilization or control of competing vegetation resulted in dramatic increases in almost every measure of productivity investigated, including height (19.7 m in the FW treatment versus 12.5 m in the C treatment at age 18 years), basal area (FW=44.2 m² ha⁻¹, F=39.6 m² ha⁻¹, W=36.6 m² ha⁻¹, C=19.9 m² ha⁻¹ at age 16 years), stemwood biomass accumulation (114 Mg ha⁻¹ in FW versus 42.8 Mg ha⁻¹ in C at age 18 years), foliar nitrogen concentration (1.53% in plots receiving fertilization versus 1.06% in unfertilized plots at age 17 years) and leaf area index (age 16-year peak projected of approximately 3.3 at age 9–10 years in F and FW plots, 2.5 in the W treatment and 1.5 in the C plots). Cultural treatments also decreased the growth ring earlywood/latewood ratio, and accelerated the juvenile wood to mature wood transition. While soil nutrient supply was a major determinant of productivity, production changes that occurred within treatments over the course of stand development were equally dramatic. For example, between age 8 and 15 years, stemwood PAI in the FW treatment declined by 275%; similarly large reductions occurred in the F and W treatments over the same time period. The reductions in PAI in the treated plots were linearly related to stand BA, suggesting the decline in productivity was associated with the onset of inter-tree competition. Responses of stemwood PAI to re-fertilization treatments at age 15 years suggests that the declines in growth and growth efficiency with time were partially attributable to nutrient limitations.     

Regional influences of soil available water-holding capacity and climate,and leaf area index on simulated loblolly pine productivity

We simulated loblolly pine (Pinus taeda L.) net canopy assimilation, using BIOMASS version 13.0, for the southeastern United States (1° latitude by 1° longitude grid cells) using a 44-year historical climate record, estimates of available water-holding capacity from a natural resource conservation soils database, and two contrasting leaf area indices (LAI) (low; peak LAI of 1.5 m² m⁻² projected, and high; 3.5 m² m⁻²). Median (50th percentile) available water-holding capacity varied from 100 to 250 mm across the forest type for a normalized 1.25 m soil profile. Climate also varied considerably (growing season precipitation ranged from 200 to 1600 mm while mean growing season temperature ranged from 13° to 26°C). Net canopy assimilation ranged from 9.3 to 19.2 Mg C ha⁻¹ a⁻¹ for high LAI and the 95th percentile of available water-holding capacity simulations.We examined the influence of soil available water-holding capacity, and annual variation in temperature and precipitation, on net canopy assimilation for three cells of similar latitude. An asymptotic, hyperbolic relationship was found between the 44-year average net canopy assimilation and soil available water-holding capacity. Shallow soils had, naturally, low water-holding capacity (<100 mm) and, subsequently, low productivity. However, median available water-holding capacity (125–150 mm) was sufficient to maintain near maximum production potential in these cells.Simulations were also conduced to examine the direct affects of soil available water on photosynthesis (P_N) and stomatal conductance (g_S) on net canopy assimilation. In the absence of water limitations on P_N and g_S, net canopy assimilation increased by only 10% or less over most of the loblolly pine region (when compared to simulations for median available water-holding capacity with water influences in place). However, the production differences between high and low LAI, at the median soil available water-holding capacity, ranged from 30% to 60% across the loblolly pine range. Vapor pressure deficit was found to dramatically reduce productivity for stands of similar LAI, incident radiation, rainfall, and available water-holding capacity. Thus, these simulations suggest that, regionally, loblolly pine productivity may be more limited by low LAI than by soil available water-holding capacity (for soils of median available water-holding capacity or greater). In addition, high atmospheric forcing for water vapor will reduce net assimilation for regions of otherwise favorable available water and LAI.     

Landscape pattern and spatial variability of leaf area index in Eastern Amazonia

Uncertainties about the implications of land-cover heterogeneity on the Amazonian carbon (C) and water cycles are, in part, related to the lack of information about spatial patterns of key variables that control these fluxes at the regional scale. Leaf area index (LAI) is one of these key variables, regulating a number of ecosystem processes (e.g. evaporation, transpiration and photosynthesis). In order to generate a sampling strategy for LAI across a section of Amazonia, we generated a landscape unit (LU) map for the Tapajós region, Eastern Amazonia, as a basis for stratification. We identified seven primary forest classes, stratified according to vegetation and/or terrain characteristics, and one secondary forest class, covering 80% of the region. Primary forest units were the most representative, covering 62% of the total area. The LAI measurements were carried out in 13 selected LUs. In each LU, we marked out three 50 m × 50 m plots giving a total number of 39 plots (9.75 ha). A pair of LAI-2000 plant canopy analysers was used to estimate LAI. We recorded a total of 25 LAI measurements within each plot. We used the field data to verify the statistical distribution of LAI samples, analyse the LAI variability within and among sites, and show the influence of sample size on LAI variation and precision. The LAI showed a high coefficient of variation at the plot level (0.25 ha), from 5.2% to 23%, but this was reduced at the landscape unit level (three co-located plots, 1.8–12%). The level of precision was <10% and 15% at the plot and landscape unit level, respectively. The LAI decreased from a dense lowland forest site (5.10) to a secondary forest (3.46) and to a pasture site (1.56). We found evidence for differences in the scale of spatial heterogeneity of closed canopy forest versus open canopy forest and palm forests. Landscape variables could, in part, explain differences in LAI among forest sites, and land use is an important modifier of LAI patterns. The stratified LAI sampling proposed in the present study could cope with three important aspects of C and water fluxes modelling: (1) optimise the information obtained from field measurements, which is an advance for models parameterisation, compared to the usual random sampling; (2) generate information for a subsequent scaling up of point field measurements to surfaces covering the whole region; and (3) build a useful basis for validation of estimations, based on remote sensing data, of LAI in the Tapajós region. The variability of LAI in the Tapajós region showed that this variable is a source of uncertainty for large-scale process modelling.     

Long term growth responses of loblolly pine to optimal nutrient and water resource availability

A factorial combination of four treatments (control (CW), optimal growing season water availability (IW), optimum nutrient availability (FW), and combined optimum water and nutrient availability (FIW)) in four replications were initiated in an 8-year-old Pinus taeda stand growing on a droughty, nutrient-poor, sandy site in Scotland County, NC and maintained for 9 years. Results for the first 4 years after treatment initiation at this study were first reported by Albaugh et al. [For. Sci. 44 (1998) 317]. The site is primarily nutrient limited and all measured stand parameters (height, basal area, leaf area index, live crown length, stem mass accumulation, current annual stem mass increment) were increased with fertilization throughout the study period. Irrigation effects were also positive for these parameters but the increases were much smaller than those found with fertilization. For example, 9 years after treatment initiation, standing stem mass was increased 100 and 25% by fertilization and irrigation, respectively, while current annual increment of stem biomass production was increased 119 and 23% by fertilization and irrigation, respectively. Interestingly, stem density (stems ha⁻¹) was not significantly affected by treatment in any year of the study. Growth efficiency (stem mass increment per unit leaf area index) was 1.9 Mg ha⁻¹ per year per LAI for CW and influenced by treatment with IW, FW, and FIW achieving growth efficiencies of 2.4, 2.7 and 2.9 Mg ha⁻¹ per year per LAI, respectively. Growth efficiency appeared to be relatively stable in the last 4 years of the study. Ring specific gravity was measured in the third, fourth, and fifth years after treatment initiation. An average reduction in ring specific gravity of 7.5% was observed with fertilization while irrigation had little effect on specific gravity in any year measured. The continuation of high growth rates with no observable growth decline in the treated stands throughout the 9-year study may be a function of the age of the stands when treatments were initiated (8 years), the very poor initial nutrient and moisture availability, and/or the application of an ongoing optimum nutrient regime at the site. The fertilized plots are now at or near an age and a size when a commercial harvest would be feasible. For the stand conditions at this site, then, the optimum nutrient availability plots have achieved high productivity throughout the economic life of the stand without measurable declines in stand productivity.     

A comparison of harvesting productivity and costs in thinning operations with and without midfield

Mechanised thinning operations can be carried out in the forest where skid roads are provided on which harvesters and forwarders can move. In the transition to continuous cover forestry (CCF) it is better to keep a thinner network of skid roads in the forest. Instead of tracks for harvesters and forwarders, these areas can be used for younger generations of trees. Moreover, fewer skid roads in the forest environment make the stand more natural. Fewer skid roads were introduced in this research as an alternative thinning operation with midfield¹ (MF) to the most popular mechanised thinning operation with skid roads² (SR). The aim of this paper is to analyse the productivity and economic aspects of thinning operations based on harvesters and forwarders, where there are different distances between skid roads. In both of the operations, harvesters and forwarders were used, but in the MF operation a chainsaw was additionally used to cut trees beyond the reach of the harvester boom. The distances between skid roads in the MF operation were 35–38 m, while in the other they were 18–20 m. The research was carried out in premature pine stands in a flat terrain in Poland. Bigger productivity and lower costs were found in the MF thinning operations. In the younger 44-year-old stand, the average harvester (Timberjack 770) productivity (in operational time) in the MF operation was 5.87 m³h⁻¹ and in the SR operation 4.52 m³h⁻¹; forwarding provided by the Vimek 606 6WD achieved a productivity of 5.03 and 4.52 m³h⁻¹, respectively. In the older 72-year-old stand, the Timberjack 1270B productivity was 11.53 m³h⁻¹ in MF and 8.70 m³h⁻¹ in SR; the Timberjack 1010B forwarder achieved 11.22 m³h⁻¹ (MF) and 8.84 m³h⁻¹(SR).The costs of harvesting and forwarding 1 m³ of wood were lower in the MF operations. In the younger stand, harvesting costs were 5.78 €/m³ (MF) and 6.72 €/m³ (SR) while forwarding costs were 1.94 and 2.18 €/m³ respectively. In the older stand, harvesting costs were 5.58 €/m³ (MF) and 6.78 €/m³ (SR); the forwarding costs were 2.65 €/m³ (MF) and 3.41 €/m³ (SR).     

Can intensive management accelerate the restoration of Brazil's Atlantic forests?

Only 7% of the once extensive forest along the eastern coast of Brazil remains, and much of that is degraded and threatened by agricultural expansion and urbanization. We wondered if methods similar to those developed to establish fast-growing Eucalyptus plantations might also work to enhance survival and growth of rainforest species on degraded pastures composed of highly competitive C₄ grasses. An 8-factor experiment was laid out to contrast the value of different intensities of cultivation, application of fertilizer and weed control on the growth and survival of a mixture of 20 rainforest species planted at two densities: 3 m × 1 m, and 3 m × 2 m. Intensive management increased seedling survival from 90% to 98%, stemwood production and leaf area index (LAI) by ～4-fold, and stemwood production per unit of light absorbed by 30%. Annual growth in stem biomass was closely related to LAI alone (r² = 0.93, p < 0.0001), and the regression improved further in combination with canopy nitrogen content (r² = 0.99, p < 0.0001). Intensive management resulted in a nearly closed forest canopy in less than 4 years, and offers a practical means to establish functional forests on abandoned agricultural land.     

Growth and carbon stocks of a spruce forest chronosequence in central Europe

Human induced changes in global environmental conditions are expected to influence or, as it is hypothesised in this study, have already influenced the biomass and growth of forest ecosystems. In this study, we reconstruct the history of tree growth and quantify the standing biomass along a chronosequence of six Norway spruce stands (Picea abies [L.] Karst; 16–142 years old) on acid soils in a mountainous region with high nitrogen deposition. The inventories of the study sites, as well as the historical stem growth of the sample trees were compared with common yield tables, representing growing conditions before 1960, to find out if and when significant changes in growth of trees had occurred. The growth at tree level (0.003–0.030 m³ yr⁻¹) was about 150–350% higher than predicted by the yield tables, independent of tree age. Because of low stand densities due to early thinning, the increase of stem growth at stand level (90% higher than yield table predictions) and the stand volume (35% higher than yield table predictions) were not as high as the increase of growth at tree level. Total biomass at stand level (including stems, branches, twigs, needles and roots) ranged between 35 and 180 t C ha⁻¹. Net primary productivity varied between 6 and 13 t C ha⁻¹ yr⁻¹. Intensive tree thinning activities probably stimulated growth of remaining trees, but the observed growth rates were beyond what would be expected from these activities exclusively. Thus it is assumed that the fertilisation effects of increased nitrogen deposition and CO₂ concentration, and improved climatic conditions due to ongoing climate change, have contributed to the observed changes in stem growth and that the thinning activities were synergetic with changing environmental conditions. The implications for carbon sinks as accountable under the Kyoto Protocol are probably small, because changes in environmental conditions are not accountable under the Kyoto Protocol and most of the observed changes in growth took place before 1990, the baseline for the Kyoto Protocol. Additionally, it is assumed that impacts on the carbon balance of forest stands due to changes in the thinning regime after 1990, which would be accountable according to article 3.4 of the Kyoto Protocol, are very small without any synergetic changes in environmental conditions.     

Production,allocation, and stemwood growth efficiency of Pinus taeda L. stands in response to 6 years of intensive management

Loblolly pine (Pinus taeda L.) is a highly plastic species with respect to growth responses to forest management. Loblolly pine is the most planted species across the southern United States, a region with the most expansive and intensively managed forest plantations in the world. Management intensity, using tools such as site preparation and fertilization, is increasing greatly in scope over time. To better define to the productive potential of loblolly pine under intensive management, the influence of 6 years of management with weed control (W), weed control plus irrigation (WI), weed control plus irrigation and fertigation (irrigation with a fertilizer solution) (WIF), or weed control plus irrigation, fertigation, and pest control (WIFP) since plantation establishment on stand productivity in loblolly pine was examined. The site is located near Bainbridge, GA (30°48′N latitude and 84°39′W longitude) and is of medium quality (site index=18 m, base age 25). Increasing management intensity greatly accelerated stand development and biomass accumulation. At age 6 total production (above plus belowground) was nearly doubled from 50 to 93 Mg ha⁻¹ in WIFP stands compared to W stands, and standing stem biomass increased from 24 Mg ha⁻¹ in W stands to 48 Mg ha⁻¹ in response to WIFP treatment. Stem current annual increment (CAI) peaked at age 5 in the WIF and WIFP stands at 17–18 Mg ha⁻¹ per year at a basal area between 18 and 21 m² ha⁻¹. Year to year variation in CAI was better explained by previous-year leaf area index (LAI) than current-year LAI. Maximum stemwood production in loblolly pine was achieved through large increases in LAI and small decreases in allocation to woody roots (tap+coarse roots) versus woody shoots (stem+branches) associated with intensive treatments.     

Feasibility of implementing thinning in even-aged <Emphasis Type="Italic">Larix olgensis</Emphasis> plantations to develop uneven-aged larch–broadleaved mixed forests

Thinning experiments were conducted in larch (Larix olgensis) plantations to assess the feasibility of converting even-aged plantation stands to uneven-aged forests with more complex stand structures. Stands established in 1965 and 1960 were thinned in 2004 (Regime A, for determining the effect of recent thinning on emergence of seedlings) and 1994 (Regime B, for examining the effects of the past thinning on establishments of recruitments), respectively, at two intensities each. Natural regeneration, together with litter depth, canopy openness and vegetation cover, was surveyed in the thinned plots. Results indicated that larch seedlings started to emerge in May, reached a peak in June, decreased from June through September, and then disappeared in October. No larch seedlings exceeded 1 year old in the thinned plots because of the low levels of light and dense litter and vegetation cover. However, there were many naturally regenerated seedlings (5–50 cm in height) and saplings (50–500 cm in height) of broadleaved tree species such as Acer spp., Fraxinus spp., Cornus controversa, Quercus mongolica, and even the climax tree species, Pinus koraiensis, in the thinned plots. The mean density of regenerated seedlings reached 6.7 and 4.5 stems m⁻² in Regimes A and B, respectively, whilst the mean density of regenerated saplings reached 4,595 stems ha⁻¹ in Regime B. These results suggest that it is impractical to turn even-aged larch plantations to uneven-aged larch forests, but it may be feasible to develop uneven-aged larch-broadleaved forests from even-aged larch plantations through thinning.     

Regeneration dynamics of Sitka spruce in artificially created forest gaps

This study examined the variation in the development of naturally regenerated and planted seedlings of Sitka spruce (Picea sitchensis (Bong.) Carr.) within gaps cut in a 32-year-old stand of the same species. The circular gaps were 20 m in diameter and designed to allow sunlight into only half of the gap floor at midsummer given the latitude of 56°45′N. Eight plots (8 m × 3 m) were laid out along a north–south transect through each gap (four within the gap and two each under the closed canopy north and south of the gap). Each plot was sub-divided and seedlings were planted into one part and the other part was left to naturally regenerate. In subsequent seasons, plots were further subdivided into ‘weed free’ and ‘vegetation left untouched’. Results showed that while the two central plots within the gaps had the highest value of canopy openness, the highest accumulated temperature and lowest soil moisture were recorded in plots that received direct sunlight. However, level of germination was significantly higher in the shaded area of the gap than in the part that received direct sunshine suggesting that higher moisture levels in shaded areas are important to successful germination. Minimal germination was recorded in the plots beneath the canopy. Seedling survival was significantly influenced by the influx of competing vegetation, but only in the part of the gaps that received direct sunlight. The success of Sitka spruce regeneration within gaps appears to depend on sufficient moisture and light to support regeneration and early growth, but not too much light to encourage the development of competing vegetation. The permanently shaded areas of the gaps appeared to offer ground conditions with sufficient moisture and light to ensure successful germination and early growth of seedlings, but without excessive competition from other vegetation.     

Logging activity and tree regeneration in an Amazonian forest

We studied the effect of experimental logging of 4 ha plots on the regeneration of tree species in a forest 90 km north of Manaus, Amazonas, Brazil. Logging resulted in a total reduction in live wood volume of 44–107 m³ ha⁻¹, although only 63% of this volume was felled, and only 43% removed from the plots. The density of established regeneration (trees and shrubs with diameter at breast height ≤10 cm, and height ≥200 cm) was greater in logged plots than in control plots when measured 3 and 7–8 years after logging. Species richness was also significantly higher in logged plots than in controls. We registered 139 species per 1000 stems, 7–8 years after logging, 143 species per 1000 stems, 3 years after logging, and 136 species per 1000 stems in control plots. Overall species composition was significantly affected by the intensity of logging damage in the plots after 7–8 years, and control plots were significantly different from plots logged 3 years previously. However, changes were not great in relation to natural variation within the forest. Most species increased in density after logging (mean=17%), and the number of individuals belonging to species with commercial value on the local market was 15% greater in logged plots than in control plots. The total potential value of the regeneration, based on the value of wood per m³ (when adult) of the individuals, was 23% higher in logged plots than in control plots, though this difference was not statistically significant. Therefore, enrichment planting is not necessary to maintain either the biodiversity, or potential economic value for wood production, of this forest.     

Effects of wood-ash on the tree growth,vegetation and substrate quality of a drained mire: a case study

The effects of wood-ash fertilisation on tree stands, soil characteristics and ground vegetation were studied on a drained pine mire in Finland (64°51′N, 26°04′E, 62 m a.s.l.). The original site type was a treeless, mesotrophic Sphagnum papillosum fen. The site was drained in 1933 and the wood-ash fertilisation experiment was started in 1947. The treatments were: (i) unfertilised, (ii) wood-ash 8 t ha⁻¹, and (iii) wood-ash 16 t ha⁻¹.Drainage and ash application had radical and long-lasting consequences on the biological activity on the site and the vegetation compartments studied. The understorey vegetation had been profoundly affected by the ash with almost complete transformation of the species and other life forms. Even 50 years after the ash treatment the changes in vegetation/site type and the tree stand were clearly visible. On the unfertilised plot, the biomass of ground vegetation consisted mostly of mosses and dwarf shrubs, but on the ash-treated plots it consisted mostly of herbs and grasses typical of upland forests.Ash treatment had promoted stem volume growth of Scots pine (Pinus sylvestris L.) substantially and for a long time. The total wood production on the ash plots during 1947–1994 was 13 and 17 times over that of the control plot. Unfertilised pine trees suffered from P and K deficiency throughout the study period. The concentrations of some plant nutrients (P, K) decreased during the past years on Ash8. No nutrient shortage afflicting the tree stand was observed on Ash16 during the study period.Ash application has also led to increased concentration of nutrients in the peat. A sizeable proportion of the mineral nutrients applied were still in the 0–20 cm peat layer. On the ash-treated plots the amount of soil nitrogen (0–20 cm) was 18 and 29 times and the amount of soil phosphorus 9 and 13 times over the amount bound by the tree stand and the ground vegetation (Ash8 and Ash16, respectively). The stock of potassium was generally small in the surface peat—only 60–90% of the amount of potassium bound in the tree stand and the ground vegetation.It was concluded that wood-ash had powerfully influenced the biological processes in surface peat. The decomposition of cellulose was significantly accelerated by both ash treatments. Ash fertilisation also increased the emissions of CO₂. The intensified decomposition rate in the litter, vegetation and peat explained to a large extent the accelerated growth of the Scots pine stands studied.     

Canopy gap characteristics and their implications for management in the temperate rainforests of southeast Alaska

In the coastal temperate rainforests of southeast Alaska, much progress has been made in describing landscape-level natural disturbances and formulating management systems that emulate those disturbances. Little is known, however, concerning canopy gaps, the dominant form of natural disturbance in the region. During June–August, 1991–1993, we characterized canopy gap patterns and dynamics at three sites in the western hemlock/blueberry/shield fern plant association in the northern portion of the Tongass National Forest.Forest area in canopy gaps ranged from 5.8 to 12.6% and averaged 8.7%. The proportion of forest area in expanded gaps ranged from 18.1 to 43.9% and averaged 27.4%. Gap and gapmaker (tree whose death or crown displacement results in the creation or expansion of a canopy gap) characteristics were generally similar among sites. The majority of canopy gaps were <50 m² in area, had a D/H ratio <0.50, were created from the death of one or two gapmakers, and had experienced gap expansion. The majority of expanded gap areas were <200 m². Gapmakers were usually snapped, had recently died (<20 years ago), and tended to be larger in diameter than surrounding overstory trees. Species composition of gapmakers was similar to surrounding overstory trees. A substantial amount of gap infilling takes place between 20 and 80 years following tree death, but gaps, or portions of gaps, can persist for >80 years. Forest turnover time was estimated to range from 230 to 920 years, and average 575 years. Canopy residence time was estimated to range between 210 and 840 years, and averaged 525 years.To emulate canopy gap dynamics in the plant association studied, forest managers should: (1) maintain a small proportion of a stand in openings within an otherwise undisturbed canopy; (2) use a combination of single tree selection and small group selection systems; (3) re-enter stands every 20–80 years; (4) select larger than average diameter crop trees in proportion to the species composition of the stand; (5) minimize soil disturbance and (6) select crop trees during re-entry so that the creation of new gaps and the expansion of old gaps is accomplished in approximately equal proportions.     

Initial impact of supervised logging and pre-logging climber cutting compared with conventional logging in a dipterocarp rainforest in Sabah,Malaysia

Pre-marked skid trails, directional felling and climber cutting when logging in tropical rainforests may be important ways of reducing damage to the forest, thus creating a healthier stand and improving future yields.This study, carried out in a virgin dipterocarp rainforest in the south of Sabah, Malaysia, compared two types of logging (both with and without pre-cutting climbers): conventional selective logging (CL) and supervised logging (SL). The latter is a selective logging system in which both pre-marked skid trails and directional felling were implemented. The pre-marked skid trails were aligned parallel to each other, spaced 62 m apart. A randomised complete block 2 × 2 factorial design was used in the experiment, consisting of 16 gross treatment plots, each of 5.76 ha with a 1 ha net plot in the centre.Fewer trees tended (0.050 < P ≤ 0.100) to be logged in SL plots than in CL plots (on average 9.4 and 13.0 trees ≥60 cm diameter breast height ha⁻¹). Pre-felling of climbers resulted in four more dipterocarp trees being logged ha⁻¹, compared with no climber cutting: a statistically significant difference (P ≤ 0.050). The basal areas lost of both large trees (≥ 60 cm dbh) and small dipterocarp trees (10–29 cm dbh) tended to differ between the logging systems, with CL leading to greater losses.There were significant differences in the residual stands left by the logging systems, with respect to the number of dipterocarps and their basal area in the diameter class 10–29 cm; ca 30% more stems being found after SL. No significant differences (or tendencies) in these variables were found in the residual stands in other diameter classes, or when trees of all species were considered.     

Forest floor vegetation plays an important role in photosynthetic production of boreal forests

We estimated gross photosynthetic production (GPP) of the forest floor vegetation in a 40-year-old Scots pine stand in southern Finland with three different methods: measurements of CO₂ exchange of single leaves of field and ground layer species, measurement campaigns of forest floor net CO₂ efflux at different irradiances with a manually operated soil chamber, and continuous measurements of forest floor net CO₂ efflux with an automatic transparent chamber system. We upscaled the measured light response curves from the manual soil chambers using the biomass distribution of the forest floor species, a modelled seasonal pattern of photosynthetic capacity and a model of light extinction down the canopy. Leaf gas exchange measurements as well as measurements of net CO₂ efflux with the manual chamber indicated saturation of photosynthesis at relatively low (50–400 μmol m⁻² s⁻¹) light levels. Leaf and patch level measurements gave similar rates of photosynthetic CO₂ fixation per unit leaf biomass suggesting that reduction in photosynthetic production due to within-patch shading was small. Upscaling of photosynthetic production to the stand level and continuous measurements with the automatic soil chambers indicated that momentary photosynthetic production by the forest floor vegetation in the summer was typically about 2 μmol m⁻² (ground) s⁻¹. Cumulative upscaled GPP over the period of no snow (from 20 April to 20 November) in year 2003 was 131 g C m⁻². Continuous measurements with the automatic soil chamber system were in line with the upscaling, the cumulative GPP being 83 g C m⁻² and the seasonal pattern of photosynthetic rate similar to that of the upscaled photosynthesis.     

Stand dynamics in a logged and silviculturally treated Costa Rican rain forest, 1988–1996

The lowland rain forests of Central America are poorly known from the standpoint of management for timber production. We studied the stand dynamics of a logged Costa Rican rain forest under three different regimes of post-logging silvicultural treatment. The site was located on low hills with Ultisols in Holdridge's Tropical Wet Forest life zone. The Pentaclethra macroloba-dominated forest had been high graded before planned management began. Management of the 540 × 540 m (29.2 ha) experimental area began with a timber harvest in the whole area during 1989–1990, 4 trees ha⁻¹ being cut overall for 10.1 m³ ha⁻¹. The experimental plots were 180 × 180 m (3.24 ha), comprising a 100 × 100 m (1.0 ha) central permanent sample plot (PSP) with a 40-m wide buffer strip. Two types of post-harvest silvicultural treatment: liberation/refinement (in 1991) and shelterwood (in 1992) were applied under a complete randomized block design with three replicates, using logged but untreated plots as controls. PSP data reported are for the 1988–1996 period for individuals with ≥10 cm DBH. The most marked changes in forest structure were caused by silvicultural treatment, basal area under the liberation/refinement treatment being reduced to ca. 65% of its probable mature forest value. Recruitment exceeded mortality in the years following intervention under all three treatments, but forest structural recovery was slowest under the liberation/refinement treatment. Post-intervention mortality rates appeared higher under the liberation/refinement treatment than under the control or shelterwood treatments, though differences were not statistically significant. In relation to tree attributes, mortality rates increased with decreasing DBH increment, crown illumination and quality of crown form. Commercial DBH increments were higher under the liberation/refinement treatment than in control plots during the 1993–1996 period. On the basis of its response to intervention during the first seven years of management, the forest appears resilient and productive; trends over time in mortality rates under the most intense silvicultural regime require close attention however. Pentaclethra-dominated forests are important components of the productive forest resources of Costa Rica and Nicaragua and, given current deforestation rates in areas such as southern Nicaragua, it is now urgent that the existing biophysical knowledge of these forests be applied to forest conservation and management.     

Dynamics of above- and below-ground biomass and nutrient accumulation in an age sequence of Nothofagus antarctica forest of Southern Patagonia

Nothofagus antarctica (Forster f.) Oersted is a deciduous tree species, which naturally grows on poorly drained or drier eastern sites in the Andes Mountain near Patagonian steppe. Above- and below-ground biomass and nutrients pools were measured in pure even-aged stands at different ages (5–220 years) and crown classes. Functions were fitted for total biomass and nutrients accumulation, and root/shoot ratio of individual trees against age. Total biomass accumulated for mature dominant trees was eight times greater than mature suppressed trees. Biomass root/shoot ratio decreased with age from 1.8 to a steady-state of 0.5. All nutrients concentration (except Ca) decreased with age and varied according to the degree of crown suppression classes. Nutrient concentrations varied between biomass pool components following the order leaves > bark > small branches > fine roots > medium roots > rooten wood > coarse roots > sapwood > heartwood. Total nutrient accumulation followed the order dominant > codominant > intermediate > suppressed trees and its accumulation rate varied over time, e.g. P accumulation rate of dominant trees increased from 0.17 g tree⁻¹ year⁻¹ during regeneration to 1.39 g tree⁻¹ year⁻¹ in mature trees. Nutrients uptake reached a peak during the period of maximum biomass production, and root/shoot ratio of nutrients decreased from its maximum value at 5 years of age (0.6, 4.0, 0.9, 1.5, 1.0 and 2.6 for N, P, K, Ca, S and Mg, respectively) to a steady-state asymptote beyond 50 years of age. Thus, accumulation of nutrients in roots was greater during the regeneration phase of stand development, and nutrient accumulation increased in above-ground over time. Also, nutrient use efficiency increased in mature trees (111–220 years) and decreased in suppressed crown classes. The equations developed for individual trees have been used to estimate stand biomass and nutrient accumulation from forest inventories data. Total stand biomass varied from 62.5 to 133.4 t ha⁻¹ and total nutrients accumulation ranged from 3 kg Mg ha⁻¹ to 1235 kg Ca ha⁻¹. Proposed equations can be used for practical purposes such as to estimate pasture nutrients requirement in a silvopastoral system based on nutrients supply from leaf litter returns, or to determine amelioration practices like debarking stems before harvesting.