Estimation of litter fall and seed production of Acacia mangium in a forest plantation in South Sumatra, Indonesia

Annual litter fall of Acacia mangium in the period of September 1995 to August 1996 was estimated at 5939 kg ha⁻¹ year⁻¹ and from September 1995 to August 1996 at 6048 kg ha⁻¹ year⁻¹, with the highest seasonal production in the dry season. The litter fall was dominated mainly by leaves, 4446 kg (75%) and 4137 kg (68%), respectively. Seed production in the litter fall was estimated at 42.4 kg ha⁻¹ year⁻¹ (4.1 million seeds ha⁻¹) and 39 kg ha⁻¹ year⁻¹ (3.8 million seeds ha⁻¹), with the highest in the dry season from June to October. The accumulated litter fall in the forest floor together with shrubs and grass provide a high fuel load, increasing fire risk.     

Biomass and carbon sequestration of ponderosa pine plantations and native cypress forests in northwest Patagonia

Fast growth tree plantations and secondary forests are considered highly efficient carbon sinks. In northwest Patagonia, more than 2 million ha of rangelands are suitable for forestry, and tree plantation or native forest restoration could largely contribute to climate change mitigation. The commonest baseline is the heavily grazed gramineous steppe of Festuca pallescens (St. Yves) Parodi. To assess the carbon sequestration potential of ponderosa pine (Pinus ponderosa (Dougl.) Laws) plantations and native cypress (Austrocedrus chilensis (Don) Flor. et Boutl.), individual above and below ground biomass models were developed, and scaled to stand level in forests between 600 and 1500 annual rainfall. To calculate the carbon sequestration baseline, the pasture biomass was simulated. Also, soil carbon at two depths was assessed in paired pine-cypress-pasture sample plots, the same as the litter carbon content of both forest types. Individual stem, foliage, branch and root log linear equations adjusted for pine and cypress trees presented similar slopes (P>0.05), although some differed in the elevations. Biomass carbon was 52.3 Mg ha⁻¹ (S.D.=30.6) for pine stands and 73.2 Mg ha⁻¹ (S.D.=95.4) for cypress forests, given stand volumes of 148.1 and 168.4 m³ ha⁻¹, respectively. Soil carbon (litter included) was 86.3 Mg ha⁻¹ (S.D.=46.5) for pine stands and 116.5 Mg ha⁻¹ (S.D.=38.5) for cypress. Root/shoot ratio was 19.5 and 11.4%, respectively. The low r/s value for cypress may account for differences in nutrient cycling and water uptake potential. At stand level, differences in foliage, taproot and soil carbon compartments were highly significative (P<0.01) between both forest types. In pine stands, both biomass and soil carbon were highly explained by the rainfall gradient (r²=0.94). Nevertheless, such a relationship was not found for cypress, possibly due to stand and soil disturbances in sample plots. The carbon baseline estimated in pasture biomass, including litter, was 2.6 Mg ha⁻¹ (S.D.=0.8). Since no differences in soil carbon were found between pasture and both forest types, additionality should be accounted only by biomass. However, the replacement of pasture by pine plantations may decrease the soil carbon storage, at least during the first years. On the other hand, the soil may be a more relevant compartment of sequestered carbon in cypress forests, and if pine plantation replaces cypress forests, soil carbon losses could cause a negative balance.     

Nitrogen-fertilization impacts on carbon sequestration and flux in managed coastal Douglas-fir stands of the Pacific Northwest

We examined whether N-fertilization and soil origin of Douglas-fir [Psuedotsuga menziesii (Mirb.) Franco] stands in western Washington state could affect C sequestration in both the tree biomass and in soils, as well as the flux of dissolved organic carbon (DOC) through the soil profile. This study utilized four forest sites that were initially established between 1972 and 1980 as part of Regional Forest Nutrition Research Project (RFNRP). Two of the soils were derived from coarse-textured glacial outwash and two from finer-textured volcanic-source material, primarily tephra, both common soil types for forestry in the region. Between 1972 and 1996 fertilized sites received either three or four additions of 224 kg N ha⁻¹ as urea (672–896 kg N ha⁻¹ total). Due to enhanced tree growth, the N-fertilized sites (161 Mg C ha⁻¹) had an average of 20% more C in the tree biomass compared to unfertilized sites (135 Mg C ha⁻¹). Overall, N-fertilized soils (260 Mg C ha⁻¹) had 48% more soil C compared to unfertilized soils (175 Mg C ha⁻¹). The finer-textured volcanic-origin soils (348 Mg C ha⁻¹) had 299% more C than glacial outwash soils (87.2 Mg C ha⁻¹), independent of N-fertilization. Soil-solution DOC collected by lysimeters also appeared to be higher in N-fertilized, upper soil horizons compared to unfertilized controls but it was unclear what fraction of the difference was lost from decomposition or contributed to deep-profile soil C by leaching and adsorption. When soil, understory vegetation and live-tree C compartments are pooled and compared by treatment, N-fertilized plots had an average of 110 Mg C ha⁻¹ more than unfertilized controls. These results indicate these sites generally responded to N-fertilization with increased C sequestration, but differences in stand and soil response to N-fertilization might be partially explained by soil origin and texture.     

Forest structure and C stocks in natural Fagus sylvatica forest in southern Europe: The effects of past management

Managed forests often differ substantially from undisturbed forests in terms of tree structure and diversity. By altering the forest structure, management may affect the C stored in biomass and soil. A survey of 58 natural stands located in the south-westernmost limit of European beech forests was carried out to assess how the C pools are affected by the changes in tree structural diversity resulting from past management. The mean tree density, basal area and the number of large trees found in unmanaged forests were similar to those corresponding to virgin beech forests in Central Europe, whereas large live trees were totally absent from partially cut stands. Analysis of the Evenness index and the Gini coefficient indicated high structural diversity in the three stand types. The results of the Kolmogorov–Smirnov test used to compare the diameter distributions of each group revealed significant differences between stand types in terms of distributions of total tree species and of Fagus sylvatica.

The mean C stocks in the whole ecosystem – trees, litter layer and mineral soil – ranged from 220 to 770 Mg ha⁻¹ (average 380 Mg ha⁻¹). Tree biomass (above and belowground), which averaged 293 Mg C ha⁻¹, constituted the main C pool of the system (50–97%). The statistical test (Kolmogorov–Smirnov) revealed differences in the distribution of C pools in tree biomass between unmanaged and partially cut stands. As a consequence of the presence of large trees, in some unmanaged stands the C stock in tree biomass was as high as 500–600 Mg C ha⁻¹. In the partially cut stands, most of the C was mainly accumulated in trees smaller than 20 cm dbh, whereas in unmanaged stands the 30% of tree C pool was found in trees larger than 50 cm dbh. Furthermore, many unmanaged stands showed a larger C pool in the litter layer. The C content of mineral soils ranged from 40 to 260 Mg C ha⁻¹ and it was especially high in umbrisols. In conclusion, the implementation of protective measures in these fragile ecosystems may help to maintain the highly heterogeneous tree structure and enhance the role of both soils and trees as long-term C sinks.     

Fluxes of CO2, CH4 and N2O from drained coniferous forests on organic soils

Fluxes of CO₂, CH₄ and N₂O were measured during two to three years at four sites, located within an area of 9 km² in southern Sweden, using dark static chamber techniques. Three of the sites were drained coniferous forests on moist organic soils that differed in forest productivity and tree species. The fourth site was an undrained tall sedge mire. Although the drained sites were all moist, with average groundwater levels between 17 and 27 cm below the soil surface, the mean annual dark forest floor CO₂ release rate was significantly higher at the drained sites, (0.9–1.9 kg m⁻² y⁻¹) than at the undrained mire site (0.8 to 1.2 kg m⁻² y⁻¹). CH₄ emissions were significantly lower from the drained sites than from the undrained mire (0.0 to 1.6 g m⁻² y⁻¹, compared to 10.6 to 12.2 g m⁻² y⁻¹), while N₂O emissions were significantly lower from the undrained site than from the drained sites (20 to 30 mg m⁻² y⁻¹, compared to 30 to 90 mg m⁻² y⁻¹). There were no clear effects of site productivity or tree species on the soil fluxes of any of the gases. The annual net primary production of the forests was modeled. All drained sites were net sinks, while the undrained mire was a net source of greenhouse gases. The estimated net greenhouse gas exchange of the drained sites was correlated with productivity: the most productive site was the largest net sink and the least productive the smallest net sink for greenhouse gases. The results indicate that, to mitigate the increase of atmospheric greenhouse gases, drained forest sites, which have been unsuccessfully drained or rewetted due to subsidence, should be managed in a way that keeps the groundwater level at a steady state.     

Burning of Amazonian forest in Ariquemes, Rondônia, Brazil: biomass, charcoal formation and burning efficiency

Biomass burning in tropical forests – the normal practice to prepare land for agriculture and ranching – has been a major source of CO₂ emitted to the atmosphere. Mass transformations by burning are still little studied in the tropics. The present study estimated parameters, such as the stock of carbon contained in the biomass, burning efficiency and the formation of charcoal and ashes in a tropical moist forest. Two sets of plots arranged in the form of ‘stars' (720 m² total) were installed in a 3.5 ha area of forest that had been felled for planting pasture at Fazenda Nova Vida, Ariquemes, Rondônia. Each ‘star' had six rays measuring 2 m × 30 m; alternating rays were designated for pre-burn and post-burn measurements. All above-ground biomass present in the plots was weighed directly before the burn in the pre-burn rays and after the burn in the post-burn rays. Pieces of wood with diameter ≥10 cm also had their biomasses estimated from volume estimates, using line-intersect sampling (LIS) in order to increase the area of sampling and to allow volume loss to be estimated as an increment based on individual pieces measured before, and after, the burn at the same point (as opposed to inferring change as a difference between independent estimates of stocks). The initial above-ground biomass (dry weight) before the burn was estimated at 306.5 ± 48.6 (mean ± SE) Mg ha⁻¹, with an additional 4.5 Mg ha⁻¹ for trees left standing. Carbon stock in the initial biomass (including trees left standing) was 141.3 (Mg C) ha⁻¹. After burning, carbon stock was reduced by 36.8% (burning efficiency). The stocks of charcoal and ash formed in the burn were, respectively, 6.4 ± 2.7 and 5.7 ± 1.0 Mg ha⁻¹. The destructive and nondestructive (LIS) methods did not differ significantly (t-test, p > 0.05) in estimating post-burn stocks of wood and charcoal. The results of this study contribute to improving the estimates of parameters needed for global carbon calculations and point to ways in which estimates of these parameters could be further improved.     

What the soil reveals: Potential total ecosystem C stores of the Pacific Northwest region, USA

How much organic C can a region naturally store in its ecosystems? How can this be determined, when land management has altered the vegetation of the landscape substantially? The answers may lie in the soil: this study synthesized the spatial distribution of soil properties derived from the state soils geographic database with empirical measurements of old-growth forest ecosystem C to yield a regional distribution of potential maximum total-ecosystem organic C stores. The region under consideration is 179,000 square kilometers extending from the southern Oregon border to the northern Washington border, and from the Pacific Ocean to the east side of the Cascade Mountains. Total ecosystem organic C (TEC) was measured in 16 diverse old-growth forests encompassing 35 stands and 79 pedons to a depth of 100 cm. The TEC ranged between 185 and 1200 Mg C ha⁻¹. On an average, 63% of TEC was in the vegetation, 13% in woody detritus, 3% in the forest floor, 7% in the 0–20 cm mineral soil, and 13% in 20–100 cm mineral soil. The TEC was strongly related to soil organic C (SOC) in the 0–20 cm mineral soil, yielding a monotonically increasing, curvilinear relation. To apply this relation to estimate the TEC distribution throughout the region, 211 map units of the state soils geographic database (STATSGO) were used. The SOC in the 0–20 cm mineral soil of the map units was consistent with values from previously measured pedons distributed throughout the region. Resampling of 13 second-growth forests 25 years after initial sampling indicated no regional change in mineral SOC, and supported the use of a static state soils map. The SOC spatial distribution combined with the quantitative old-growth TEC–SOC relation yielded an estimate of potential TEC storage throughout the region under the hypothetical condition of old-growth forest coverage. The area-weighted TEC was 760 Mg C ha⁻¹. This is 100 Mg C ha⁻¹ more than a previous estimate based on a coarser resolution of six physiographic provinces, and 400 Mg C ha⁻¹ more than current regional stores. The map of potential TEC may be useful in forecasting regional C dynamics and in land-management decisions related to C sequestration.     

Effects of harvesting intensity on carbon stocks in eastern Canadian red spruce (Picea rubens) forests: An exploratory analysis using the CBM-CFS3 simulation model

Carbon stocks and stock changes in a chronosequence of 24 red spruce (Picea rubens Sarg.) dominated stands in Nova Scotia, Canada, were compared against predictions from the Carbon Budget Model of the Canadian Forest Sector (CBM-CFS3). Regression analysis of the observed versus simulated total ecosystem C stocks indicates the model's predictions accounted for 81.1% of the variation in the observed biomass data and for 63.2% of the variation in total ecosystem C data; however, the simultaneous F-test for bias was significant. Discrepancy between the observed and simulated total ecosystem C data was primarily caused by differences in dead organic matter C pool estimates, with the model consistently predicting higher soil C throughout stand development. Changes to model parameters were not warranted however, as the field data measured only a portion of the mineral soil profile represented in the model.

Clear-cut and partial-cut harvesting scenarios for red spruce stands were simulated to examine the impacts of clear-cut and partial-cut harvesting on C stocks. Total ecosystem C increased in the partial-cut stand throughout the 240-year simulation from 308.9 to 327.3 Mg C ha⁻¹, while it decreased in the clear-cut stand to 305.8 Mg C ha⁻¹. Enhanced C sequestration in the partial-cut stand was a consequence of the residual standing biomass providing a continuous source of litterfall and reducing decomposition rates of the forest floor. Choice of harvest system clearly affects forest ecosystem C stocks, but also affects the amount of C removed from forests to meet society's needs. Over the period of the simulation, partial cutting provided 115.6 Mg C ha⁻¹ of merchantable timber, while clear-cutting provided 132.4 Mg C ha⁻¹. Strategies aimed at using forest management to mitigate atmospheric C increases need to assess both the direct impacts on the forest ecosystem and the indirect impacts through product and energy substitution associated with the use and storage of harvested biomass.     

Growth, biomass production and nutrient accumulation by seven tree species irrigated with municipal effluent at Wodonga, Australia

In the Murray-Darling basin, irrigation of tree crops is being evaluated as an alternative method for the disposal of municipal effluent. A study was carried out at Wodonga in which seven tree species were irrigated with effluent for a period of 4 years. Irrigation was calculated weekly on the basis of pan evaporation and rainfall during the preceding week. Annual irrigation varied between 1190 mm and 1750 mm with a total input over the 4-year-period of 4940 mm.

Height and diameter growth varied significantly between species. At age 4, mean dominant height of Eucalyptus grandis, E. saligna and Populus deltoides × P. nigra ranged from 14.3 to 15.0 m compared with 6.6 to 9.8 m for Casuarina cunninghamiana, E. camaldulensis, P. deltoides and Pinus radiata. Wood production of the faster-growing species (E. grandis and E. saligna) was approximately 130 m³ ha⁻¹, or around 32 m³ ha⁻¹ year⁻¹ over a 4-year period. This was nearly three-fold the production of the other native species and twice that of Pi. radiata. Volume growth of P. deltoides × P. nigra (85 m³ ha⁻¹) was significantly greater than that of P. deltoides (42 m³ ha⁻¹).

Accumulation of nutrients in the above-ground biomass varied significantly between species and ranged from 24 to 41 g m⁻² for N, 2.6 to 5.9 g m⁻² for P, 0.5 to 9.2 g m⁻² for Na, 12 to 27 g m⁻² for K, 7 to 52 g m⁻² for Ca and 3.1 to 7.9 g m⁻² for Mg. Nutrient accumulation was generally greater in species with a comparatively large crown biomass relative to stem size such as C. cunninghamiana and E. camadulensis. Average nutrient accumulation by trees as a percentage of input from effluent was estimated at 19% for N, 9% for P, 1% for Na, 14% for K, 52% for Ca and 32% for Mg.

Results of this study indicate the importance of selecting species on the basis of not only growth but also nutrient accumulation to optimise renovation of wastewater by tree plantations.     

Tree regeneration responses to microsite characteristics following a severe tornado in the Georgia Piedmont, USA

For 3 years following a severe, November 1992 tornado, abundance and growth of tree regeneration in intact forest-floor and windthrow-pit microsites were studied in three mixed pine and hardwood stands in the Georgia Piedmont, USA. The research had two objectives: (1) determine if performance of tree regeneration differed between microsite types and between pre- and post-tornado cohorts on intact forest-floor microsites, and (2) determine if variation in light and soil water availability from the disturbance affected performance of eight species artificially seeded into intact forest-floor microsites. Near each of the 42 sample points (12–20 per site) spaced on 15 m grids, species and height of tree seedlings were recorded within a 1 m radius plot of intact forest floor and the nearest windthrow pit. Intact forest-floor microsites were dominated by two late-successional species, Acer rubrum (3.5 pre-tornado stems per m² and 1.8 post-tornado stems per m²) and Ostrya virginiana (2.6 post-tornado stems per m²), while windthrow pits were dominated by an early successional species, Liriodendron tulipifera (1.7 stems per m²). Although seedling survival did not vary significantly among species or microsite types, first-year height of seedlings in intact forest-floor microsites (24 cm) was significantly greater than those in windthrow-pit microsites (19 cm). Second-year height growth of new seedlings of Cornus florida in intact forest-floor microsites (52 cm) significantly exceeded that of many other combinations of species and microsite type. Species artificially-seeded into intact forest-floor microsites in 1994 and 1995 varied considerably in emergence (<1–24%), survival (<1–16%), and height (5–15 cm), and those with the heaviest seed survived best. From 1994 to 1995, average gap fraction, an index of light availability, decreased 17% from 0.23 to 0.19. Soil water content in 1995 under gap fractions of greater than 0.3 (21%) averaged significantly less than under gap fractions of either 0.1–0.3 (24%) or less than 0.1 (26%). Significant positive correlations were detected for relationships of emergence and survival of several species to soil water content. Overall rankings of seedling performance between microsites (intact forest floor>windthrow pits), between cohorts (pre-tornado>post-tornado origin), and among species (mid- to late->early-successional species) indicate that advanced regeneration and new seedlings of A. rubrum, O. virginiana, and C. florida will be long-term dominants of the understory because of their high abundance, initial growth responses, and shade tolerance.     

Structure, allometry, and biomass of plantation Metasequoia glyptostroboides in Japan

We quantified structural features and the aboveground biomass of the deciduous conifer, Metasequoia glyptostroboides (Hu and Cheng) in six plantations in central Japan. In order to derive biomass estimates we dissected 14 M. glyptostroboides trees into three structural components (stem wood, branch wood and foliage) to develop allometric equations relating the mass of these components and of the whole tree to diameter at breast height (DBH). We found robust relationships at the branch and whole tree level that allow accurate prediction of component and whole tree biomass. Dominant tree height was similar within five older (>40 years) plantations (27–33 m) and shorter in a 20-year-old plantation (18 m). Average stem diameter varied from 12.8 cm in the youngest stand to greater than 35 cm in the oldest stand.

Metasequoia have relatively compact crowns distributed over the top 30% of the tree although the youngest stand had the deepest crown relative to tree height (up to 38%). At the individual tree level in older stands, 87% of the aboveground biomass was allocated to the stem, 9% to branch wood and 4% to foliage. We found little difference in the relative distribution of above ground biomass among the stands with the exception of lower foliage biomass in larger diameter trees. Total aboveground biomass of the older stands varied twofold, ranging from a maximum of 450 Mg ha⁻¹ in a 42-year-old stand to a minimum of 196 Mg ha⁻¹ in a 48-year-old stand. Total above ground biomass of the 20-year-old stand was 176 Mg ha⁻¹.     

Long-term soil potassium availability from a Kanhapludult to an aggrading loblolly pine ecosystem

A long-term (1962 to 1990) forest biogeochemistry study in the southeastern Piedmont of the USA provided estimates of soil K release in response to forest regrowth. We investigated the sources of soil K that buffered the exchangeable K pools during forest growth and we estimated soil K release rates through greenhouse and acid extraction studies for comparison to our field estimate.

In these acid Kanhapludults, derived from granitic-gneiss, the disparity between measured depletions of soil exchangeable K and estimated forest removals indicated a buffering of exchangeable K on the order of 0.31 kmol_c ha⁻¹ per year. Non-exchangeable K extracted by boiling with 1 M HNO₃ exceeded exchangeable K by up to 40-fold. Non-exchangeable K was not depleted during the three decades of stand growth, however, thus was not the long-term source of exchangeable K buffering. Total K in these soils ranged from 0.4 to 3.8% by weight. Mineralogical data indicated a presence of hydroxy-interlayered vermiculite throughout the upper 4 m of soil for <2 μm clay fraction and a presence of micaceous minerals in the 2 to 45 μm silt fraction. XRD analysis of micaceous flakes extracted from 4 to 8 m in the soil indicated a presence of muscovite mica.

Estimated K releases in the greenhouse and extraction studies were generally consistent with long-term results. The accumulation of K during two rotations of pine seedling growth in the greenhouse exceeded the measured depletions in exchangeable and non-exchangeable K over all soil depths tested by 0.007 to 0.026 cmol_c kg⁻¹. Potassium removal by sequential extraction/incubations with 1 mM HCl and 1 mM oxalic acid continued through 24 extractions and K recovered in extract solutions exceeded the sum of depletions in exchangeable and non-exchangeable K pools by 0.001 to 0.028 cmol_c kg⁻¹. These excess removals in plant uptake or solution recovery indicate a release of mineral K. Thirty-day extractions with H⁺-resins in both 1 mM HCl and 1 mM oxalic acid were well fit by the Elovich equation but were not well correlated with plant K uptake in the greenhouse study. The release rate coefficients ranged from 0.012 to 0.025(cmol_c kg⁻¹) h⁻¹.

Extrapolations to annual releases of K in the greenhouse and sequential extraction studies were a similar order of magnitude as long-term releases estimated at the long-term Calhoun plots. Surface soil (0 to 15 cm) releases ranged from 0.15 to 0.65 kmol_c ha⁻¹per year while deeper soils ranged up to 1.54 kmol_c ha⁻¹per year. Results indicate that soils similar to those at Calhoun that contain a similar micaceous and HIV component will be able to supply K at rates adequate to keep pace with demands of forest regrowth even under intensive forest management.     

Microsite patterns of conifer seedling establishment and growth in a mixed stand in the southern Alps

In many forests of the Alps, permanent forest cover and, therefore, its continuous renewal is the main silvicultural goal. Regenerating these forests must be based on a sound understanding of the ecology of the tree species in question. The regeneration of silver fir (Abies alba Mill.) and Norway spruce (Picea abies (L.) Karst.) in the upper montane zone of the southern central Alps has so far received little attention. The aim of this study was, therefore, to characterize the ecological niche for the establishment and growth of silver fir and Norway spruce seedlings in this zone. The study was conducted as a case study on a plot of 1.5 ha in the upper montane zone of southern Switzerland, at an altitude of 1380 m. The basal area of the mixed Norway spruce–silver fir stand was 43.7 m² ha⁻¹. Seedlings were censused on 375 plots of 1.0 m² area. Seedling density was 1.30 ± 4.25 m⁻² (mean ± standard deviation) for Norway spruce and 0.87 ± 1.50 m⁻² for silver fir. Logistic regression models were used to test the effect of microsite characteristics on seedling occurrence, and general linear models for effects on seedling height growth and biomass increment. Most seedlings received less than 10% light (photosynthetic photon flux density) as compared to values in the open. Silver fir occurrence was positively related to microsites at the edge of canopy trees, but unrelated to ground cover type, light and micro-relief. Norway spruce occurrence was only, and positively, related to the presence of mosses. Height growth and biomass increment of seedlings of both species were only loosely correlated with microsite conditions. Seedlings without canopy cover grew faster than those under canopy cover, probably as a result of light and moisture limitations under the canopy of adult trees. Diffuse radiation was positively correlated with average annual biomass increment of silver fir, but not of Norway spruce seedlings. In general, the results suggest that silver fir seedlings have less specific microsite requirements than Norway spruce seedlings in terms of ground cover. They are also more shade tolerant, and therefore, grow faster than Norway spruce in low-light environments of the upper montane zone of the southern Alps. While the microsite concept can be helpful in designing silvicultural operations, it has limitations when only patch characteristics that are easy to assess are used, and others neglected. Finally, our study suggests that counting the number of green shoots is a promising method to quickly and non-destructively estimate the biomass of a great number of small seedlings.     

Biomass distribution and productivity ofPinus edulis—Juniperus monosperma woodlands of north-central Arizona

Above-ground biomass distribution, leaf area, above-ground net primary productivity and foliage characteristics were determined for 90- and 350-year-oldPinus edulis-Juniperus monosperma ecosystems on the Colorado Plateau of northern Arizona. These ecosystems have low biomass, leaf area and primary productivity compared with forests in wetter environments. Biomass of the 350-year-old pinyon-juniper stand examined in this study was 54.1 mg ha⁻¹; that of the 90-year-old stand was 23.7 mg ha⁻¹. Above-ground net primary production averaged 2.12 mg ha⁻¹ year⁻¹ for the young and 2.88 mg ha⁻¹ year⁻¹ for the mature stand; tree production was about 80% of these values for both stands. Projected ecosystem leaf area (LAI) of the stands was 1.72 m² m⁻² and 1.85 m² m⁻², respectively. Production efficiency (dry matter production per unit leaf area) was 0.129 kg m⁻² year⁻¹ for the young, and 0.160 kg m⁻² year⁻¹ for the mature stand. Production efficiency of the study sites was below the 0.188 kg m⁻² year⁻¹ reported for xeric, pure juniper stands in the northern Great Basin. Biomass of pinyon-juniper ecosystems of northern Arizona is generally below the 60–121 mg ha⁻¹ reported for pinyon-juniper stands of the western Great Basin in Nevada. A climatic gradient with summer precipitation decreasing between southeast Arizona and northwest Nevada occurs in the pinyon-juniper region. Great Basin pinyon-juniper ecosystems lie at the dry-summer end of this gradient while pinyon-juniper ecosystems of the Colorado Plateau lie at about the middle of this gradient. In spite of wetter summers, pinyon-juniper ecosystems of northern Arizona are less productive than those of the Great Basin.     

Some nutrient dynamics associated with litterfall and litter decomposition in hoop pine plantations of southeast Queensland, Australia

Litterfall was collected over a 12-month period with littertraps in hoop pine (Araucaria cunninghamii) plantations aged 10, 14 and 62 years in southeast Queensland, Australia. The bulk of litterfall occurred during spring, mainly as hoop pine foliage with the annual litterfall ranging between 6.0 and 10.9 t ha⁻¹, respectively, for the younger stands (10 and 14 years) and the mature 62-year old stand. The amount of nitrogen (N) and phosphorous (P) recycled annually through litterfall was lower in the younger stands (28–37 kg N ha⁻¹ and 4.4–5.3 kg P ha⁻¹) compared with that of the mature stand (85 N ha⁻¹ and 6.2 kg P ha⁻¹). The N and P retranslocated during senescence varied across the three stands studied with a trend for N and P retranslocation to increase as availability of soil mineral-N decreased.

Decomposition of the hoop pine foliage component of litter was also studied in the same stands using a litterbag technique and mass-balance analysis. The estimated half-life of hoop pine foliage mass ranged between 1.5 and 1.8 years. Litter-mass loss was strongly correlated with litter substrate quality indicators of N, C, P, C/N ratio, lignin, lignin/N ratio and polyphenols. During the course of the study, there was no difference in litter-mass loss between the stands of different ages. During the 15-month period, the order of element release from the hoop pine litter was K>Na>C>Mg>P, with N, Ca and Mn generally demonstrating varying degrees of net accumulation. During the course of the study, the lignin/C ratio of the hoop pine litter increased from 0.61 to 0.96. This suggested that the litter-C was predominantly in a recalcitrant form and, therefore, the associated N was unlikely to be rapidly released in the hoop pine litter layer.     

Required sample size for estimating soil respiration rates in large areas of two tropical forests and of two types of plantation in Malaysia

We estimated the required sample sizes for estimating large-scale soil respiration (for areas from 1 to 2 ha) in four ecosystems (primary and secondary forests, and oil palm and rubber plantations) in Malaysia. The soil respiration rates were 769 ± 329 mg CO₂ m⁻² h⁻¹ in the primary forest (2 ha, 50 sample points), 708 ± 300 mg CO₂ m⁻² h⁻¹ in the secondary forest (2 ha, 50 points), 815 ± 363 mg CO₂ m⁻² h⁻¹ in the oil palm plantation (1 ha, 25 points), and 450 ± 178 mg CO₂ m⁻² h⁻¹ in the rubber plantation (1 ha, 25 points). According to our sample size analysis, the number of measurement points required to determine the mean soil respiration rate at each site with an error in the mean of no more than 10% ranged from 67 to 85 at the 95% probability level. These results suggest that evaluating the spatial heterogeneity of soil respiration rates in the tropics may require more measurement points than in temperate forests.     

Pre-commercial thinning effects on growth, yield and mortality in even-aged paper birch stands in British Columbia

The aim of this study was to quantify 5-year growth, yield and mortality responses of 9- to 13-year-old naturally regenerated, even-aged paper birch (Betula papyrifera Marsh.) stands to pre-commercial thinning in interior British Columbia. The study included four residual densities (9902–21,807 stems ha⁻¹ (unthinned control), 3000, 1000 and 400 stems ha⁻¹) and four sites with 3-fold within-site replication in a randomised block design. The largest, straightest, undamaged trees were selected to leave during thinning. Thinning reduced stand basal area from 5.90 m² ha⁻¹ in the control to 2.50, 1.53 and 0.85 m² ha⁻¹ in the three thinning treatments, representing 42, 26 and 15% of control basal area, respectively. After 5 years, total stand volume per plot remained lower in the three thinning treatments than the control (50.20, 30.07, 18.99 and 11.86 m³ in the control, 3000, 1000 and 400 stems ha⁻¹ treatments), whereas mean stand diameter, diameter increment, height, and height increment were increased by thinning, and top height (tallest 100 trees ha⁻¹) was unaffected. When a select group of crop trees (largest 250 trees ha⁻¹) in the thinning treatments was compared with the equivalent group in the control, there was a significant increase in mean diameter, diameter increment, basal area, basal area increment, and volume increment. Mean height, height increment, top height, and total volume were unaffected by thinning. Crop tree diameter increment was the greatest following thinning to 400 stems ha⁻¹ for all diameter classes. Thinning to 1000 stems ha⁻¹ resulted in lower diameter increment than thinning to 400 stems ha⁻¹ but tended to have higher volume increment. Dominant trees responded similarly to subdominant trees at 400 stems ha⁻¹, but showed the greatest response at 3000 stems ha⁻¹. Results suggest that pre-commercial thinning of 9–13-year-old stands to 1000 stems ha⁻¹ would improve growth of individual trees without seriously under-utilising site resources.     

Woody vegetation structure and composition along a protection gradient in a miombo ecosystem of western Tanzania

Fully protected areas such as National Parks are often assumed to be the best way to conserve plant diversity and maintain intact forest composition and structure. To evaluate this assertion, we sampled trees in areas with four different levels of protection: a National Park, where the protection level is very high, a Game Controlled Area which allows tourist hunting of big game animals, a Forest Reserve which allows selective harvest of trees, and an Open Area where human populations have unrestricted access to forest resources. All four land management areas had healthy size-class distributions with greater numbers of juvenile trees (2–10 cm DBH) than adults. Surprisingly, mean stem density of trees was highest (947 stems ha⁻¹) in the Game Controlled Area but was lowest (635 stems ha⁻¹) in the National Park. The former had the highest basal area value (24 m² ha⁻¹) while the human-inhabited Open Area had the lowest (11 m² ha⁻¹). Species richness in the Forest Reserve and Game Controlled Area was significantly higher than in the other areas. The total number of plots with unique species not found anywhere else was lowest in the National. Our measures of forest structure and composition therefore show that fully protected National Parks do not necessarily conserve the greatest diversity of tree species or unique species, indicating that a suite of different types of protection strategies may be the key to conservation of trees in these African dry tropical forests.     

Litterfall, litter decomposition and nutrient dynamics in a subtropical natural oak forest and managed plantation in northeastern India

In northeastern India, subtropical forests are over-exploited for timber, fuel wood and common agricultural practice like shifting cultivation, which are responsible for the degradation of natural forest. In degraded areas, large-scale plantations of different species of Quercus have been raised since 1980 for the production of economic Tasar silk. Conversion of natural forest into plantation affects the process of nutrient cycling due to management practices. Thus, it would be of importance to study the litterfall, litter decomposition process and the factors regulating the rate of litter decay in these ecosystems to improve recommendations for their management and conservation. We recorded litterfall by using litter traps and decomposition of leaf litter by nylon net bag technique to understand the amount of organic matter and nutrient return and their release in soils of forest and plantation in Manipur, northeast India. Total litterfall was 419.9 g m⁻² year⁻¹ in plantation and 547.7 g m⁻² year⁻¹ in forest. Litter decomposition rate was faster at plantation site than the forest in the early stage of litter decomposition whereas the reverse was observed at later stages of decomposition. Stepwise regression analysis showed the significant role of relative humidity and mean temperature on mass loss rates in the forest. Relative humidity, maximum temperature, population of fungi and actinomycetes were the best predictor variables for mass loss rates in plantation. Nutrient retranslocation efficiency and the immobilization of N and P in forest litter were higher than plantation. This suggests that Q. serrata growing in natural ecosystem in oligotrophic condition adapted strong nutrient conservation mechanisms to compete with the other plant species for the meager soil nutrients. The same species in plantation loses these adaptive capabilities because of exogenous supply of nutrients and in the absence of intense competition with other plant species. Thus, the optimization of organic and chemical fertilizer input in plantation is recommended for maintaining the soil fertility level to produce quality leaf for silkworm by conserving essential nutrients in the system.     

Biomass production and allocation, including fine-root turnover, and annual N uptake in lysimeter-grown basket willows

Annual net primary production (NPP) and N uptake were estimated for lysimeter-grown basket willows (Salix viminalis L.) during 3 years after planting. The willows were grown in a stand structure and continuously supplied with water and liquid fertilizer through drip tubes. The lysimeters contained either clay from the site or washed quartz sand. Shoot growth and leaf litter were measured and fine-root dynamics observed in minirhizotrons. Destructive samples were taken annually in late autumn and entire root systems were washed out. Dry mass and N content of all plant parts were determined. Fine-root production was estimated by two methods, based on destructive samplings and observations in minirhizotrons.

The proportion of biomass allocated below ground increased considerably when estimates based on accumulated NPP were compared with those based on standing dry mass. In the first year, 49 and 58% of annual NPP in willows grown in clay and sand, respectively, was belowground. In subsequent years the proportions were 36–38% and 33–40%. Most belowground production was fine roots. Relatively more N was used belowground in the first year than subsequently, but no substrate-induced differences were observed in the allocation pattern. Both annual NPP and N uptake was always higher in plants in clay than in those in sand: in the final 2 years, 21–22 tonnes DM ha⁻¹ year⁻¹ and 190 kg N ha⁻¹ year⁻¹ in clay, and 9–10 tonnes DM ha⁻¹ year⁻¹ and 100 kg N ha⁻¹ year⁻¹ in sand.