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.     

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.     

Effects of uneven-aged silviculture on the stand structure,species composition,and economic returns of loblolly pine stands

Uneven-aged silviculture in loblolly pine (Pinus taeda L.) stands has many economic and ecological benefits. Here, the consequences of various uneven-aged management regimes are predicted with the SouthPro simulator. Results indicate that target distributions for pines with residual merchantable basal areas of ≈12.5 m² ha⁻¹, maximum diameters of ca. 40 cm, and q-ratios of 1.2–1.25 for 2.5 cm DBH classes are likely to provide high economic returns on good sites when combined with hardwood control. Increasing this maximum diameter would enhance tree-size diversity, but reduce sawtimber production and profits. Retaining a hardwood component with 1.15–2.3 m² ha⁻¹ of basal area could enhance tree-species diversity, but this too would result in moderate reductions in income. Insisting on maximizing tree-size diversity or tree-species diversity among softwoods, soft hardwood, and hard hardwoods would be quite costly in terms of lost income and production. Results also illustrate how short-term economic incentives can lead to high-grading practices, despite substantial reductions in stand productivity and net returns in the long term.     

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.     

Necromass in undisturbed and logged forests in the Brazilian Amazon

Necromass is an important stock of carbon in tropical forests. We estimated volume, density, and mass of fallen and standing necromass in undisturbed and selectively logged forests at Juruena, Mato Grosso, Brazil (10.48°S, 58.47°W). We also measured standing dead trees at the Tapajos National Forest, Para, Brazil (3.08°S, 54.94°W) complementing our earlier study there on fallen necromass. We compared forest that was selectively logged using reduced-impact logging methods and undisturbed forest. We estimated necromass density accounting for void volume for necromass greater than 10 cm diameter at Juruena for five decay classes that ranged from freshly fallen (class 1) to highly decayed material (class 5). Average necromass density adjusted for void space (±S.E.) was 0.71 (0.02), 0.69 (0.04), 0.60 (0.04), 0.59 (0.06), and 0.33 (0.05) Mg m⁻³ for classes 1 through 5, respectively. Small (2–5 cm) and medium (5–10 cm) size classes had densities of 0.52 (0.02) and 0.50 (0.04) Mg m⁻³, respectively. The average dry mass (±S.E.) of fallen necromass at Juruena was 44.9 (0.2) and 67.0 (10.1) Mg ha⁻¹ for duplicate undisturbed and reduced impact logging sites, respectively. Small and medium sized material together accounted for 12–21% of the total fallen necromass at Juruena. At Juruena, the average mass of standing dead was 5.3 (1.0) Mg ha⁻¹ for undisturbed forest and 8.8 (2.3) Mg ha⁻¹ for forest logged with reduced impact methods. At Tapajos, standing dead average mass was 7.7 (2.0) Mg ha⁻¹ for undisturbed forest and 12.9 (4.6) Mg ha⁻¹ for logged forest. The proportion of standing dead to total fallen necromass was 12–17%. Even with reduced impact harvest management, logged forests had approximately 50% more total necromass than undisturbed forests.     

Carbon and nutrients loos in aboveground biomass along a fire induced forest-savanna gradient in the Gran Sabana,southern Venezuela

Forest degradation and savannization are critical environmental issues associated with forest fires in the Gran Sabana, southern Venezuela. Yet little is known about the ecological consequences resulting from the conversion of forest to savanna in this region. In this study we quantified the change in C and nutrients in aboveground biomass along a fire induced gradient consisting of unburned tall primary forest (TF), slightly fire-affected medium forest (MF), strongly fire-affected low forest (LF) and savanna (S). Total aboveground biomass (TAGB) decreased from 411 Mg ha⁻¹ in TF to 313 Mg ha⁻¹ in MF, 13 Mg ha⁻¹ in LF and 5 Mg ha⁻¹ in S. The pools of C and nutrients in TAGB decreased 13–25% from TF to MF, 88–97% from TF to LF and 97–98% from TF to S. In TF and MF, about 40% of C and over 80% of base cations (Ca, K and Mg) was stored in TAGB, whereas the bulk of N and P were stored in the soil (90% of N and 72% of P). This distribution of elements was different in LF and S, where about 50% of base cations were stored in TAGB, and more than 94% of C, 98% of N and 87% of P were stored in the mineral soil. The large amount of elements stored in the biomass of the tall unburned forest demonstrates the high sensitivity of this ecosystem to fire. The change from tall forest to low forest and savanna implies large losses of C and nutrients stored in aboveground biomass and soils (namely 390–399 Mg C ha⁻¹, 11–13 Mg N ha⁻¹, 70–72 kg P ha⁻¹, 783–818 kg K ha⁻¹, 736–889 kg Ca ha⁻¹, and 200–225 kg Mg ha⁻¹). Such drain of C and nutrients in soils extremely low in silicates, which can replenish the lost nutrients by weathering reduces the recuperation chance of these ecosystems and therefore their future capacity to sequester C and accumulate nutrients.     

Estimation of biomass and net primary productivity of major planted forests in China based on forest inventory data

Reforestation and afforestation have been suggested as an important land use management in mitigating the increase in atmospheric CO₂ concentration under Kyoto Protocol of UN Framework Convention on climate change. Forest inventory data (FID) are important resources for understanding the dynamics of forest biomass, net primary productivity (NPP) and carbon cycling at landscape and regional scales. In this study, more than 300 data sets of biomass, volume, NPP and stand age for five planted forest types in China (Larix, Pinus tabulaeformis, Pinus massoniana, Cunninghamia lanceolata, Pouulus) from literatures were synthesized to develop regression equations between biomass and volume, and between NPP and biomass, and stand age. Based on the fourth FID (1989–1993), biomass and NPP of five planted forest types in China were estimated. The results showed that total biomass and total NPP of the five types of forest plantations were 2.81 Pg (1 Pg = 10¹⁵ g) and 235.65 Mg ha⁻¹ yr⁻¹ (1 Mg = 10⁶ g), respectively. The area-weighted mean biomass density (biomass) and NPP of different forest types varied from 44.43 (P. massoniana) to 146.05 Mg ha⁻¹ (P. tabulaeformis) and from 4.41 (P. massoniana) to 7.33 Mg ha⁻¹ yr⁻¹ (Populus), respectively. The biomass and NPP of the five planted forest types were not distributed evenly across different regions in China. Larix forests have the greatest variations in biomass and NPP, ranging from 2.7 to 135.37 Mg ha⁻¹ and 0.9 to 10.3 Mg ha⁻¹ yr⁻¹, respectively. However, biomass and NPP of Populus forests in different region varied less and they were approximately 50 Mg ha⁻¹ and 7–8 Mg ha⁻¹ yr⁻¹, respectively. The distribution pattern of biomass and NPP of different forest types closely related with stand ages and regions. The study provided not only with an estimation biomass and NPP of major planted forests in China but also with a useful methodology for estimating forest carbon storage at regional and global levels.     

Effect of complete competition control and annual fertilization on stem growth and canopy relations for a chronosequence of loblolly pine plantations in the lower coastal plain of Georgia

Stem growth, developmental patterns and canopy relations were measured in a chronosequence of intensively managed loblolly pine stands. The study was located on two distinct sites in the lower coastal plain of Georgia, USA and contained a factorial arrangement of complete control of interspecific competition (W) and annual nitrogen fertilization (F). The W treatment increased growth rate for several years, while the F treatments led to sustained growth increases. The combination of the W and F treatments resulted in more than 180 Mg ha⁻¹ stem biomass production at age 15 which is more than double the production of control treatment. Stem biomass production is continuing to increase through age 15 as indicated by the current annual increment in stem biomass continuing to exceed the mean annual increment in stem biomass. The F treatment decreased wood quality by decreasing whole tree latewood specific gravity from 0.565 to 0.535 and by lengthening the transition from juvenile to mature wood from 4 to 5 years. Increased rates of stem growth in response to cultural treatments were largely mediated by increased leaf area, with strong functional relationships between leaf area index and current annual increment. However, growth efficiency (stem production per unit of leaf area) decreased with stand age. These results indicate that nutrient amendments are necessary for sustaining high rates of stand development on relatively nutrient poor lower coastal plain soils.     

Spatial distribution of biomass in forests of the eastern USA

We produced a map of the biomass density and pools, at the county scale of resolution, of all forests of the eastern US using new approaches for converting inventoried wood volume to estimates of above and belowground biomass. Maps provide a visual representation of the pattern of forest biomass densities and pools over space that are useful for forest managers and decision makers, and as databases for verification of vegetation models. We estimated biomass density and pools at the county level from the USDA Forest Service, Forest Inventory and Analysis database on growing stock volume by forest type and stand size-class, and mapped the results in a geographic information system. We converted stand volume to aboveground biomass with regression equations for biomass expansion factors (BEF; ratio of aboveground biomass density of all living trees to merchantable volume) versus stand volume. Belowground biomass was estimated as a function of aboveground biomass with regression equations. Total biomass density for hardwood forests ranged from 36 to 344 Mg ha⁻¹, with an area-weighted mean of 159 Mg ha⁻¹. About 50% of all counties had hardwood forests with biomass densities between 125 and 175 Mg ha⁻¹. For softwood forests, biomass density ranged from 2 to 346 Mg ha⁻¹, with an area-weighted mean of 110 Mg ha⁻¹. Biomass densities were generally lower for softwoods than for hardwoods; ca. 40% of all counties had softwood forests with biomass densities between 75 and 125 Mg ha⁻¹. Highest amounts of forest biomass were located in the Northern Lake states, mountain areas of the Mid-Atlantic states, and parts of New England, and lowest amounts in the Midwest states. The total biomass for all eastern forests for the late 1980s was estimated at 20.5 Pg, 80% of which was in hardwood forests.     

Biomass and nutrient accumulation in pure and mixed plantations of indigenous tree species grown on poor soils in the humid tropics of Costa Rica

Aboveground biomass and nutrients and soil chemical characteristics were examined in young plantations of four indigenous tree species: Hieronyma alchorneoides, Vochysia ferruginea, Pithecellobium elegans, and Genipa americana, growing in mixed and pure stands at La Selva Biological Station, Costa Rica. Total tree biomass production rates ranged from about 5.2 Mg ha⁻¹ year⁻¹ for G. americana to 10.3 Mg ha⁻¹ year⁻¹ for H. alchorneoides pure stands, and for the species mixture it was about 8.9 Mg ha⁻¹ year⁻¹. Branches and foliage formed 25–35% of total tree biomass but they represented about 50% of total tree nutrients. H. alchorneoides, the four species mixture, and P. elegans had the greatest accumulations of total aboveground nutrients per hectare. The importance of the plantation floor as a nutrient compartment varied temporally. When forest floor litter biomass was at its peak, plantation floor litter N, Ca, and Mg were roughly equal to, or greater than stem nutrients for all species except for P. elegans. For P. elegans, the plantation floor consistently represented a very low proportion of total aboveground nutrients. G. americana and V. ferruginea trees showed 55–60% less biomass accumulation in mixed than in pure stands while H. alchorneoides and P. elegans trees grew 40–50% more rapidly in mixture. P. elegans foliage had 60% lower Ca but higher P concentrations in mixed than in pure stands, and G. americana had higher foliar Mg in mixed than in pure stands. V. ferruginea stands had the highest concentrations of soil Ca, Mg, and organic matter, particularly in the top layers. Relative to pure plantations, soil nutrient concentrations in mixed plantations were intermediate for N, P, and K, but lower for Ca and Mg. The results of this study can be used in the selection of tree species and harvest designs to favor productivity and nutrient conservation.     

Profitability potential for Pinus taeda L. (loblolly pine) short-rotation bioenergy plantings in the southern USA

The use of renewable resources is important to the developing bioenergy economy and short rotation woody crops (SRWC) are key renewable feedstocks. A necessary step in advancing SRWC is defining regions suitable for SRWC commercial activities and assessing the relative economic viability among suitable regions. The goal of this study was to assess the potential profitability, based on obtainable yield and economic feasibility; of Pinus taeda L. (loblolly pine) across 13 states of the southern USA. A process-based growth model, 3PG, produced estimated yields of P. taeda in terms of mean annual increment (MAI) that were evaluated as internal rate of return on investment (IRR) and land expectation value (LEV). Coastal areas (southeast Texas, southwest Louisiana, and northern Florida) have the highest potential MAI production ranging from 13.7 to 18.9 Mg ha^− 1 yr^− 1. LEVs ranged from − 1126 to 3111 $ ha^− 1 on upland sites and − 2261 to 2341 $ ha^− 1 on lowland sites. IRR ranged from − 0.3% to 14.2% on uplands and − 2.9% to 10.4% on lowlands. On soils of the same textural class, LEV and IRR were higher on uplands relative to lowlands given lower site preparation costs, although the projected yield from upland soils are generally lower than those from lowland soils. The highest LEV and IRR were in northern Florida, southern Alabama, southern Georgia, and southern South Carolina. The lowest LEV and IRR were in Virginia and northern North Carolina. Spatially categorizing suitable lands in biological and economic terms can use geographic information system technology to advantage in combination with societal considerations to begin to answer sustainability questions as well as identify suitable sites for bioenergy plantations.     

Carbon sources and sinks in high-elevation spruce–fir forests of the Southeastern US

This paper examines carbon (C) pools, fluxes, and net ecosystem balance for a high-elevation red spruce–Fraser fir forest [Picea rubens Sarg./Abies fraseri (Pursh.) Poir.] in the Great Smoky Mountains National Park (GSMNP), based on measurements in fifty-four 20 m × 20 m permanent plots located between 1525 and 1970 m elevation. Forest floor and mineral soil C was determined from destructive sampling of the O horizon and incremental soil cores (to a depth of 50 cm) in each plot. Overstory C pools and net C sequestration in live trees was estimated from periodic inventories between 1993 and 2003. The CO₂ release from standing and downed wood was based on biomass and C concentration estimates and published decomposition constants by decay class and species. Soil respiration was measured in situ between 2002 and 2004 in a subset of eight plots along an elevation gradient. Litterfall was collected from a total of 16 plots over a 2–5-year period.The forest contained on average 403 Mg C ha⁻¹, almost half of which stored belowground. Live trees, predominantly spruce, represented a large but highly variable C pool (mean: 126 Mg C ha⁻¹, CV = 39%); while dead wood (61 Mg C ha⁻¹), mostly fir, accounted for as much as 15% of total ecosystem C. The 10-year mean C sequestration in living trees was 2700 kg C ha⁻¹ year⁻¹, but increased from 2180 kg C ha⁻¹ year⁻¹ in 1993–1998 to 3110 kg C ha⁻¹ year⁻¹ in 1998–2003, especially at higher elevations. Dead wood also increased during that period, releasing on average 1600 kg C ha⁻¹ year⁻¹. Estimated net soil C efflux ranged between 1000 and 1450 kg C ha⁻¹ year⁻¹, depending on the calculation of total belowground C allocation. Based on current flux estimates, this old-growth system was close to C neutral.     

Productivity,nutrient cycling,and succession in single- and mixed-species plantations of Casuarina equisetifolia,Eucalyptus robusta,and Leucaena leucocephala in Puerto Rico

Tree growth, biomass productivity, litterfall mass and nutrient content, changes in soil chemical properties and understory forest succession were evaluated over a 8.5-year period in single- and mixed-species (50 : 50) plantations of two N₂-fixing species, Casuarina equisetifolia and Leucaena leucocephala, and a non-fixing species, Eucalyptus robusta. At the optimal harvest age for maximum biomass production (4 years), total aboveground biomass ranged from 63 Mg ha⁻¹ in the Eucalyptus monoculture to 124 Mg ha⁻¹ in the Casuarina/Leucaena mixture, and was generally greater in the mixed-species than in single-species treatments due to increased productivity of the N-fixing species in the mixed stands. Total litterfall varied from 5.3 to 10.0 Mg ha⁻¹ year⁻¹ among treatments, or between 5.9% and 13.2% of net primary production. Litterfall production and rates of nutrient return for N, P, K, Ca and Mg were generally highest for Leucaena, intermediate for Casuarina and lowest for Eucalyptus. These rates were usually higher in the mixed-species than in monospecific stands due to differences in biomass productivity, but varied considerably depending on their species composition. Total system carbon and nutrient pools (in biomass plus soils to 40-cm depth) for N, P, K, Ca, Mg, Mn at four years were consistently greater in the plantation treatments than in the unplanted control plots. Relative to the single-species plantations, these system pools were generally larger in the mixed-species plantations for C (−10% to +10%), N (+17% to +50%), P (−1% to +63%), K (−19% to +46%), Ca (−10% to +48%), Mg (+5% to +57%) and Mn (+19% to +86%). Whole-tree harvests at four years would result in substantial system carbon and nutrient losses, although these estimated losses would not exceed the estimated gains realized during the four-year period of tree growth at this site. At 7.5 years, soil organic matter and effective cation exchange capacity were reduced in all plantation treatments relative to the control. Changes in soil nutrient content from 0 to 7.5 years were highly variable and not significantly different among treatments, although stands containing Leucaena generally showed higher rates of nitrogen and phosphorus accretion in soils than those with Eucalyptus and/or Casuarina. Natural regeneration of secondary forest tree and shrub species increased over time in all plantation treatments. A total of 24 native or naturalized forest species were recorded in the plantations at 8.5 years. Woody species abundance at this age was significantly greater beneath Casuarina than either Eucalyptus or the Eucalyptus/Leucaena mixed stands. Species richness and diversity, however, were greatest beneath stands containing Eucalyptus and/or Leucaena than in stands with Casuarina.     

Biomass,harvestable area,and forest structure estimated from commercial timber inventories and remotely sensed imagery in southern Amazonia

The purpose of this study was to determine if spatially-explicit commercial timber inventories (CTI) could be used in conjunction with satellite imagery to improve timber assessments and forest biomass estimates in Amazonia. As part of a CTI, all commercial trees ≥45 cm DBH were measured and georeferenced in 3500 ha of a logging concession in NW Mato Grosso, Brazil. A scientific inventory was conducted of all trees and palms ≥10 cm DBH in 11.1 ha of this area. A total of >20,000 trees were sampled for both inventories. To characterize vegetation radiance and topographic features, regional LANDSAT TM and ASTER images were obtained. Using a stream network derived from the ASTER-based 30 m digital elevation model (DEM), a procedure was developed to predict areas excluded from logging based on reduced impact logging (RIL) criteria. A topographic index (TI) computed from the DEM was used to identify areas with similar hydrologic regimes and to distinguish upland and lowland areas. Some timber species were associated with convergent landscape positions (i.e., higher TI values). There were significant differences in timber density and aboveground biomass (AGB) in upland (6.0 stems ha⁻¹, 33 Mg ha⁻¹) versus lowland (5.4 stems ha⁻¹, 29 Mg ha⁻¹) areas. Upland and lowland, and timber and non-timber areas could be distinguished through single and principal component analysis of LANDSAT bands. However, radiance differences between areas with and without commercial timber on a sub-hectare scale were small, indicating LANDSAT images would have limited utility for assessing commercial timber distribution at this scale. Assuming a 50 m stream buffer, areas protected from logging ranged from 7% (third order streams and above) to 28% (first order and above) of the total area. There was a strong positive relationship between AGB based on the scientific inventory of all trees and from the commercial timber, indicating that the CTI could be used in conjunction with limited additional sampling to predict total AGB (276 Mg ha⁻¹). The methods developed in this study could be useful for facilitating commercial inventory practices, understanding the relationship of tree species distribution to landscape features, and improving the novel use of CTIs to estimate AGB.     

Nitrogen budgets for Scots pine and Norway spruce ecosystems 12 and 7 years after the end of long-term fertilisation

The magnitude of nitrogen storage and its temporal change in forest ecosystems are important when analysing global change. For example, the accelerated growth of European forests has been linked to increased nitrogen deposition, but the changes in the N inputs that cause long-term changes in ecosystems have not yet been identified. We used two Swedish forest optimum nutrition experiments with Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies (L.) Karst.) to study the long-term fate of N applied to these forest ecosystems. In the pine experiment, in addition to fertiliser (NPK) application, soil acidity was manipulated by application of lime and dilute sulphuric acid. From the spruce experiment, we selected treatments with similar fertiliser doses as in the pine experiment and with and without lime addition.We quantified various terms in the N budget 12 years (pine) and 7 years (spruce) after the last N addition. In the pine stand the NPK-treatment was the only treatment to produce a significant increase in N in the tree biomass (97% above control), whereas in the spruce stand the N additions increased tree N in all treatment combinations (207% above control). In the pine stand the relative distribution of nitrogen between trees and soil did not vary across treatments, with trees containing around 12% of ecosystem N and humus containing around 44% of soil N. The increases in N stocks in the pine stands were mainly in the soil. In contrast, in the spruce ecosystem trees accumulated most of the added N and the increase in the soil was restricted to the humus layer.In the pine ecosystem, large losses of added N (between 254 and 738 kg ha⁻¹ out of 1040 kg ha⁻¹ added as fertiliser) occurred, whereas in the spruce ecosystem we recovered more N than could be accounted for by inputs (between 250 and 591 kg ha⁻¹). There was no clear pattern in the interaction between acidification/liming and N additions.     

Effects of experimental acidification,nitrogen addition and liming on ground vegetation in a mature stand of Norway spruce (Picea abies (L.) Karst.) in SE Sweden

During the period 1976–1991, a combined experiment of acidification, liming and nitrogen addition in a mature spruce stand was conducted at Farabol in south-east Sweden. The aim of this study was to investigate the effects of these treatments on the ground vegetation 0, 1, 5 and 15 years after experimental establishment. The treatment regimes were nitrogen (200 kg N ha⁻¹, repeated three times at 4–5-year intervals, totally 600 kg N ha⁻¹), sulphur powder (50 and 100 kg S ha⁻¹ a⁻¹, totally 600 and 1200 kg ha⁻¹), sulphur plus nitrogen (600+600 kg ha⁻¹) and limestone (500 kg ha⁻¹ a⁻¹, i.e. totally 6000 kg ha⁻¹). The results showed that nitrogen addition and liming promoted the abundance of the grass Deschampsia flexuosa, while acidification had a negative effect on D. flexuosa and herbs in the field layer. There was a negative reaction giving immediate damage to the bryophytes in connection with additions of nitrogen, sulphur powder and lime. The magnitude of damage and the capacity to recover varied among species as well as among treatments. The recovery from immediate damage after liming was much faster than after the treatments with sulphur powder and/or nitrogen. A negative interaction between sulphur powder and nitrogen was found for herbs and mosses where the combined effects were stronger than the effects of a single treatment alone. Acidification also had a negative effect on the total number of species. The results of this study showed that acidification and nitrogen deposition could negatively influence forest vegetation by changing the nutrient availability in the soils. Liming led to an improved growth of the forest ground vegetation and the flora changed towards a more nitrophilic species composition.     

New management options in chestnut coppices: an evaluation on ecological bases

The effects of long rotation periods and heavy low thinning on chestnut (Castanea sativa Mill.) coppices have been evaluated from a bio-ecological point of view. Aboveground tree biomass and its partitioning, bio-ecological indexes such as litter production, leaf area index (LAI), radiation regime, and growth efficiency have been analyzed for 4 years in thinned and unthinned permanent plots established in a coppice stand aged 11 years under normal rotation (NR) and in a coppice stand aged 35 years under long rotation (LR). A decrease in LAI, litter production and growth rate with the age was observed. LR showed high current annual increments (>16.0 m³ ha⁻¹ per year and 8.0 Mg ha⁻¹ per year for volume and aboveground biomass, respectively). Only slight differences in growth efficiency were observed. The adoption of heavy thinning (one third of basal area removal) affected stocking and determined significant differences in the light regime below the canopy, amount of gaps in canopy cover and LAI values, particularly in the years immediately after thinning, whereas slight differences were observed in growth efficiency. Nevertheless, chestnut showed a good aptitude, more evident in the younger stand, to re-build a homogeneous canopy cover: only a few years after thinning, canopy cover characteristics of thinned plots were similar to those of control plots and differences were not significant. Growth and increments in thinned plots were practically the same as in control plots, a consequence of consistently higher performances of released trees in the thinned plots. The results concur to a positive evaluation of a cultivation system based on long rotation periods and heavy thinnings, not only for the obtainable revenue, but especially from a bio-ecological point of view and make it a valid alternative either to abandonment or traditional over-exploitation of chestnut coppice stands.     

Genotype × environment interactions in selected loblolly and slash pine plantations in the Southeastern United States

Few studies have quantified the combined effects of silvicultural treatments and genetic improvement on unit area production of full-sib family blocks of loblolly (Pinus taeda L.) and slash pine (P. elliottii Engelm. var. elliotttii). Efficient operational deployment of genetic materials requires an understanding of possible site and silvicultural treatment interactions to maximize yield potential. We examined genotype (family) by environmental interactions (G × E) through age 5 years using a factorial experiment consisting of silvicultural treatment intensity (operational versus intensive), planting density (1334 versus 2990 trees ha⁻¹) and families (seven elite full-sib loblolly and six elite full-sib slash pine families). In January of 2000, randomized complete block, split-plot experiments were installed at two locations for each species in southeast Georgia and northeast Florida. Five years after planting, both loblolly and slash pine demonstrated significant interactions among several factors: genotype × location (p < 0.028 and p < 0.016, respectively), genotype × silvicultural treatment intensity (p < 0.055 and p < 0.059), and silvicultural treatment intensity × density (p < 0.002 and p < 0.001) for basal area (BA) and standing stem volume (VOL). Genotype × silvicultural treatment interactions were positive, with the best overall performing families responding the greatest to intensive treatment. There were changes in slash pine family rankings between locations, which were partly explained by reductions in growth associated with a combination of fusiform rust infection [Cronartium quercum (Berk.) Miyabe ex Shirai f. sp. fusiforme] and wind damage from the 2004 hurricane season. No three-way interactions, which included family, were evident and all genetic sources were stable across the contrasting planting densities. At age 5, loblolly pine outperformed slash pine (p < 0.0001), especially under the intensive silvicultural intensity. While loblolly performance was similar whether deployed in mixtures or pure family blocks, slash pine tended to be more productive in intimate mixtures than when grown in pure family blocks (p = 0.0754 for aboveground biomass).