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.     

Nitrogen deposition in Casuarina equisetifolia (Forst.) plantation stands in the dry tropics of Sonbhadra, India

Periodic variations in the concentration, deposition and canopy impact of different forms of N on annual N deposition through rainfall, throughfall and stemflow in 5 and 8 year old stands of Casuarina equisetifolia were studied. Throughfall and stemflow ranged from 70 to 76% and 5–6% of annual precipitation respectively. The total N deposition by rainfall was 11.1 kg ha⁻¹ year⁻¹, and by throughfall was 13.6 kg ha⁻¹ year⁻¹ and 16.5 kg ha⁻¹ year⁻¹ in 5-year-old and 8-year old plantations, respectively. The quantities of N deposited through stemflow in the two plantations were nearly identical, accounting for 1.6 kg ha⁻¹ year⁻¹. Observations of the monthly deposition of NH₄,N, NO₃-N, Kjeldahl-N and organic-N revealed that maximum deposition occurred in July and the minimum in September. Organic-N deposition was 17% less (5-year) than the rainwater content. Net deposition of N, as an effect of canopy, was 7–8.7 kg ha⁻¹ year⁻¹, which was added directly to the available nutrient pool of soil.     

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.     

Comparison of field methods for measuring soil respiration: a static alkali absorption method and two dynamic closed chamber methods

The objectives of the present study were to compare the static alkali absorption (AA) and dynamic closed chamber (DC) methods for measuring soil respiration, and to evaluate the effects of methodological differences on estimating annual mean soil respiration rate in a natural forest. For the AA method, we used Kirita’s method [Jpn. J. Ecol. 21 (1971) 119] using an alkali-soaked sponge disc that covers nearly the same area as that covered by a chamber. For the DC method, we used both the LI-6200 system (DC-62 method) and the newer LI-6400 system (DC-64 method) (LI-COR, Lincoln, NE, USA). Comparative measurements were conducted on five occasions during the study period (November 1998–October 1999) at a Quercus serrata forest in Japan. Daily mean soil respiration rates obtained by the AA, DC-62 and DC-64 methods for a 24 h period were in the ranges 205–578, 147–629 and 165–734 mg CO₂ m⁻² h⁻¹, respectively. The daily mean soil respiration rates obtained by the AA method were 79–128% of those obtained by the DC-64 method. When the daily mean soil respiration rate obtained by the DC-64 method was below 300 mg CO₂ m⁻² h⁻¹, the daily mean soil respiration rate obtained by the AA method was an average of 26% higher than that obtained by the DC-64 method. When the daily mean soil respiration rate obtained by the DC-64 method was above 300 mg CO₂ m⁻² h⁻¹, the daily mean soil respiration rate obtained by the AA method was an average of 19% lower than that obtained by the DC-64 method. However, at the present site, there was a little difference between the two methods as for estimating annual mean soil respiration rate, and therefore the AA method improved by Kirita [Jpn. J. Ecol. 21 (1971) 119] is suggested to be a useful method for estimating annual mean soil respiration in the forest. The daily mean soil respiration rates obtained by the DC-62 method were systematically 10–24% lower (an average of 15% lower) than those obtained with the DC-64 method, and the annual mean rate was lower than that estimated by the AA method.     

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.     

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.     

Hybrid and clonal variability of nutrient content and nutrient use efficiency in Eucalyptus stands in Congo

For 20 years, there has been 42,000 ha estate of clonal Eucalyptus plantations around Pointe-Noire in Congo on sandy soils that have very low reserves of available nutrients. These plantations have been based on a natural hybrid (E. PF1). This hybrid is being replaced by E. urophylla × E. grandis (UG), a more productive hybrid developed by the breeding program of UR2PI. A study of biogeochemical cycles showed that nutrient removal by harvesting is the main nutrient output in the E. PF1 ecosystem. It is therefore important to quantify the nutrient content (NC) in both hybrids to compare corresponding nutrient removal values.

The work dealt with four UG clones and the most planted clone of E. PF1. Twelve trees per clone were sampled at the logging age (8 years) in a clonal test for UG clones and in a nearby stand for E. PF1. Tables were established to predict, from girth at breast height (C_1.30 m), the biomass and nutrient content of stemwood, bark, dead and living branches, leaves, and were applied to the inventory of the different stands to evaluate corresponding biomass, NC and nutrient use efficiency (NUE) on a per-hectare basis.

Total biomass differed between the two hybrids and among UG clones: 109 t ha⁻¹ for E. PF1 and 108–155 t ha⁻¹ for UG clones. In E. PF1 trees, total NC was globally lower for N, K, and Mg, but greater for P and Ca. In stemwood, nitrogen content was similar for both hybrids. By contrast, in UG clones, NC was much lower for P (−72%) and Ca (−40% to −55%). The same trends were observed for NUE: equivalent for both hybrids for N, but higher in UG clones for P (+72%) and Ca (+43% to +59%). A marked variability among clones was observed for K and Mg. UG clones allocated proportionally more nutrients in leaves than E. PF1.

These results show that clones should not be selected only on growth traits but also on NUE and on the concentration of nutrients in tree components removed by harvesting. It will be then possible to limit the cost of fertilising needed to maintain stand growth and soil fertility.     

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.     

Irradiance in thinned Norway spruce (Picea abies) stands and the possibilities to prevent suckers of broadleaved trees

Both incoming shortwave radiation (R_g) and photosynthetically active radiation (PAR) in percentage of full daylight were measured at the same time by point and strip sampling in four plots (0.1 ha) of Picea abies (L.) Karst. The standard deviations (%) of R_g and PAR were, respectively, 11.1 and 9.8 at 64 points, 15.7 and 13.9 at 32 points, and 24.7 and 23.8 at 16 points per plot.

A period of at least 40 s per strip (30 m min⁻¹) gives a CV (coefficient of variation) of 30%. There is no significant difference between relative irradiance (RI) estimated by the point method (64 points) and by the strip method (8 strips). Curves of RI (R_g and PAR) and basal area (m² ha⁻¹), diameter sum (m ha⁻¹) and density (stems ha⁻¹) of fifteen trials with different thinning programmes are presented. Irradiance (R_g) in heavily thinned stands was 3–14% of irradiance on an open place. The irradiance, R_g, in extra-heavily thinned stands is 12–27%, and in unthinned stands, 1–3% that of an open place. The R_g curve lies above the PAR curve in all cases. Some practical implications of the study are presented. Heavy thinning of Norway spruce stands gives RI (R_g) values 10% at basal area of 25m² ha⁻¹ which is necessary to minimize development of suckers of broadleaved trees.     

Effect of clear-cutting and site preparation on the level and quality of groundwater in some headwater catchments in eastern Finland

The impact of forest management (clear-cutting and site preparation) on stream hydrology has been studied in five small catchments in eastern Finland from 1991 and on groundwater levels and quality from 1994. The period 1992–1996 was a calibration period and in the autumn of 1996, 10% and 30% of the area of two of these catchments were clear-cut according to the forest management plan. Regeneration was carried out by disc-plowing in the autumn of 1998 and planting with Scots pine (Pinus sylvestris) and Norway spruce (Picea abies) seedlings in the spring of 1999. The depth and quality of groundwater was monitored with four to nine groundwater wells installed in each catchment. There were 32 wells in all, 16 on upland mineral soils and 16 on peatlands (the perforated part of the pipe was totally put into the underlying mineral soil at eight sites). Sampling was made monthly during spring (March–May) and autumn (November–December) and bimonthly during summer (June–October). The samples were chemically analyzed for pH, electrical conductivity, and concentrations of total P and Fe (before filtration), and total N, NO₃⁻-N, NH₄⁺-N, total P, PO₄³⁻-P, SO₄²⁻-S, Ca²⁺, Mg²⁺, K⁺, Na⁺, Mn, Zn, Fe, Al³⁺, Cl⁻ (after filtration through 0.45 μm membrane filter). Data collected until the end of 2001 are reported. Groundwater was found in the down-slope wells in lower-lying areas, but not in those installed on the slopes with a thin (1–2 m) soil layer. Clear-cutting did not significantly affect groundwater levels in the wells. Nitrate N concentrations increased from 0.03 mg L⁻¹ level after clear-cutting and again after site preparation in the wells on upland soils and peatlands receiving water from the managed parts of the catchment. In one well at the lower edge of the managed area NO₃⁻-N concentrations reached 1–1.4 mg L⁻¹ in 2001 (fifth year after clear-cutting, third after disc-plowing), but mean concentrations remained <0.3 mg L⁻¹. Chloride concentrations also increased (50–100%) after treatments but the concentrations of other solutes showed no significant effect of treatment. It was concluded that changes in groundwater quality and quantity related to the clear-cutting were small and did not represent a danger to water quality or quantity.     

Seedling emergence, growth, and allocation of Oriental bittersweet: effects of seed input, seed bank, and forest floor litter

The establishment of invasive plant populations is controlled by seed input, survival in the soil seed bank, and effects of soil surface disturbance on emergence, growth, and survival. We studied the invasive vine Celastrus orbiculatus Thunb. (Oriental bittersweet) to determine if seedlings in forest understory germinate from the seed bank or from seed rain. We also conducted a greenhouse experiment to investigate the role of leaf litter mass and physical texture on seedling survival, growth, and allocation. In the understory of an invaded mixed hardwood forest, we measured seed input, seedling emergence with seed rain, and seedling emergence without seed rain. Mean seed rain was 168 seeds m⁻²: mean seedling emergence was 107 m⁻², and there was a strong correlation between seed rain and seedling emergence. The ratio of seedlings to seed input (0.61) was close to the seed viability (0.66) leaving very few seeds to enter the seed bank. Seed bank germination under field conditions was low (1 seedling m⁻²). Soil cores were incubated in a greenhouse to determine seed bank viability, and germination from these soil cores did not occur. To determine how litter affects seedling establishment and growth, we measured seedling emergence and biomass allocation in a greenhouse experiment. Seeds were placed below intact and fragmented deciduous leaf litter in amounts ranging from zero to the equivalent of 16 Mg ha⁻¹. Seedling emergence was not affected by fragmented litter, but decreased to <20% as intact litter increased to 16 Mg ha⁻¹. Increasing litter resulted in greater allocation to hypocotyl and less to cotyledon and radicle, and this effect was greater in intact litter. C. orbiculatus seedlings achieve emergence through forest floor litter through plasticity in allocation to hypocotyl growth. The low survival of C. orbiculatus in the seed bank suggests that eradication of seedling advance regeneration and adult plants prior to seed rain may be an effective control strategy. However, the intact forest floor litter of an undisturbed forest will not prevent seedling establishment.     

Effects of light quality on gas exchange and dry matter partitioning in Eucalyptus grandis W. Hill ex Maiden

Stockplants of Eucalyptus grandis were pruned to a height of 7–10 cm and after 3 weeks were placed in growth cabinets set at a photon flux density (PFD) of 200 μmol m⁻² s⁻¹ and red to far-red ratios of 0.4, 0.7, 1.3, 3.5 or 6.5. Experiments tested the effects of light quality on growth and gas exchange of stockplants. Light quality did not affect the total shoot dry weight (DW), root DW or shoot to root ratio of stockplants or their total leaf area. However, there were significant effects of light quality on: (i) plant height, which was greatest at red:far-red (R:FR) ratios of 0.4 and 0.7; (ii) partitioning of DW between leaves and stems, with greater stem DW and less leaf DW at low R:FR ratios (0.4 and 0.7); (iii) partitioning of DW and leaf area between the most dominant shoot and all other (non-dominant) shoots; (iv) specific leaf area, which was greatest at low R:FR ratios. In the above characters, the dominance ratio (ratio of most dominant shoot to sum of all other shoots) was greatest at low R:FR ratios and least at ratios of 3.5 and 6.5. Photosynthetic rate per unit leaf area and leaf chlorophyll concentration significantly increased with increasing R:FR ratio. However, photosynthesis per unit chlorophyll concentration was significantly greater at low R:FR ratios. Generally, light quality had no significant effect on photosynthetic rate per leaf or per unit dry weight, but rates of transpiration, stomatal conductance and water use efficiency increased with an increase in R:FR ratio. These data indicate that compensatory changes in plant morphology and gas exchange caused equality in total dry weight per plant between treatments. The above effects of light quality on dry matter partitioning and gas exchange had important effects on the size, number, morphology and physiology of subsequently collected cuttings for vegetative propagation.     

Bud activity of decapitated, nursery-grown plants ofTriplochiton scleroxylon in Nigeria: effects of light, temperature and humidity

This study is a continuation of earlier work on apical dominance inTriplochiton scleroxylon K. Schum., aimed at the development of an early clonal selection test for branching habit and yield in tree improvement programmes. Decapitated plants of five to seven clones were treated with either: (i) two photon flux densities (photosynthetically active radiation at full or 10% sunlight, approximately 2000 and 200 μmol quanta m⁻²s⁻¹ measured at midday on a clear day); (ii) two temperatures (ambient 30°C or 40°C in a polythene tent); (iii) two humidities (ambient=40–70% relative humidity at 30°C or with humidification=95–100% relative humidity at 28°C). Bud activity, shoot length and numbers of leaves produced following decapitation were all increased by high photon flux density and high humidity. In contrast to similar studies under glasshouse conditions in Britain, dominance was not re-established following the initial period of sprouting. The sprouting phase was similar in Nigeria and Britain. Clonal differences in bud activity were similar under different treatments, although there were minor changes in ranking. Increasing the air temperatures from 30 to 40°C resulted in leaf areas of 165 cm² and 23 cm², respectively. Normal polarity of lateral shoot production following decapitation was completely inverted by high temperature.     

Dynamics and potentials of carbon sequestration in managed stands and wood products in Finland under changing climatic conditions

Carbon (C) sequestration was studied in managed boreal forest stands and in wood products under current and changing climate in Finland. The C flows were simulated with a gap-type forest model interfaced with a wood product model. Sites in the simulations represented medium fertile southern and northern Finland sites, and stands were pure Scots pine and Norway spruce stands or mixtures of silver and pubescent birch.

Changing climate increased C sequestration clearly in northern Finland, but in southern Finland sequestration even decreased. Temperature is currently the major factor limiting tree growth in northern Finland. In southern Finland, the total average C balance over the 150 year period increased slightly in Scots pine stands and wood products, from 0.78 Mg C ha⁻¹ per year to 0.84 Mg C ha⁻¹ per year, while in birch stands and wood products the increase was larger, from 0.64 Mg C ha⁻¹ per year to 0.92 Mg C ha⁻¹ per year. In Norway spruce stands and wood products, the total average balance decreased substantially, from 0.96 Mg C ha⁻¹ per year to 0.32 Mg C ha⁻¹ per year. In northern Finland, the total average C balance of the 150 year period increased under changing climate, regardless of tree species: in Scots pine stands and wood products from 1.10 Mg C ha⁻¹ per year to 1.42 Mg C ha⁻¹ per year, in Norway spruce stands and wood products from 0.69 Mg C ha⁻¹ per year to 0.99 Mg C ha⁻¹ per year, and in birch stands and wood products from 0.43 Mg C ha⁻¹ per year to 0.60 Mg C ha⁻¹ per year.

C sequestration in unmanaged stands was larger than in managed systems, regardless of climate. However, wood products should be included in C sequestration assessments since 12–55% of the total 45–214 Mg C ha⁻¹ after 150 years' simulation was in products, depending on tree species, climate and location. The largest C flow from managed system back into the atmosphere was from litter, 36–47% of the total flow, from vegetation 22–32%, from soil organic matter 25–30%. Emissions from the production process and burning of discarded products were 1–6% of the total flow, and emissions from landfills less than 1%.     

Regeneration of timber trees in a logged tropical forest in North Bolivia

For sustainable forest management, it is important to know the response of timber species to the change in environment caused by logging. We performed a 2-year study on germination, survival and growth of four timber species, Cedrela odorata, Swietenia macrophylla, Hymenaea courbaril, and Cariniana micrantha, and one non-commercial species Tachigali vasquezii. We sowed seeds of these species in five microenvironments: log landing, gap-crown and gap-trunk, skidder trail and understory, in a tropical lowland moist rain forest in northern Bolivia. We related seed and seedling performance to light availability, soil compaction, and plant competition. Germination did not differ significantly between microenvironments but survival of germinated seeds for most species was significantly higher (P < 0.05) in the log landing (46–100%) than in the understory (0–7%). After 2 years, the tallest plants were always found in the log landing (119–190 cm) and the smallest in the understory (12–26 cm) caused by a higher relative height growth rate (RHGR) in the log landing (0.003–0.004 cm cm⁻¹ per day) compared to the understory (0.000–0.001 cm cm⁻¹ per day). During the first year RHGR was positively related to canopy openness for all species and negatively to the number of overtopping competitors for three species. During the second year also water infiltration explained observed variation to RHGR. These results show that abandoned log landings and logging gaps are suitable environments for the regeneration of timber species studied. This finding suggests that the removal of competitors in log landings and logging gaps combined with leaving seed trees near these microenvironments or sowing seeds, will improve regeneration of timber species in tropical forests.     

Productivity and economics of mixed two-storied spruce and birch stands in Southern Finland simulated with empirical models

The profitability of growing a naturally emerged birch (Betula pendula Roth or Betula pubescens Ehrh.) overstory in a young Norway spruce (Picea abies (L.) Karst.) plantation was examined with empirical stand structure, growth and yield, logging cost, and logging damage models. In the projected alternatives, an overstory of either birch species was thinned to 200–1000 stems per ha at the age of 15 years and retained for another 15 years. Development of the remaining spruce stand was simulated up to rotation age (70–85 years). Alternative treatments included removing the overstory completely at 15 years, and managing a pure spruce stand that was kept free of birch throughout.

Growing a birch overstory of 200–1000 stems per ha up to age 30 years resulted in a 61–93 m³ ha⁻¹ or 9.1–16.8% yield loss for the spruce stand due to growth retardation, and a mortality of 382–498 out of 1900 stems per ha through logging damage. This was compensated for or exceeded by the additional yield of the birch (54–173 m³ ha⁻¹) except for the lowest stocking (200–400 stems per ha) alternatives with B. pubescens. Treatment regimes with a birch overstory were clearly the most profitable alternatives, yielding up to 151% (B. pendula) and 113% (B. pubescens) of the net present value of the pure spruce alternative, at 4% interest rate. Removing the birch overstory already at 15 years was the least profitable alternative with 79 and 83% net present values, respectively. The most profitable treatment with current technology, price, and cost structure appears to be to grow 500–800 birch per ha up to about age 40 years for B. pendula and 45–50 years for B. pubescens.     

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.     

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.     

Use of a gradient of N-deposition to calculate effect-related soil and vegetation measures in deciduous forests

Deposition of N and S has increased since the 1950s in most European countries and N accumulates in ecosystems that are not N saturated. This study shows long-term effects of a (modelled) N deposition of 7–17 kg N ha⁻¹ per year on biological and chemical processes in soil, vegetation composition, and functional types of field-layer plant species in deciduous forests. Soil pH largely determined the response of the soil processes, emphasising the importance to compare soils of similar acidity regarding the effects of N deposition. The most pronounced effects were demonstrated for the most acid study plots. When we compared regions with a deposition of 7 and 17 kg N ha⁻¹ per year we found a 40–80% higher soil N mineralisation rate, 2–90% higher nitrification rate and 10–25% lower C:N ratio in the region with the highest deposition. Similar but smaller differences were indicated when regions with a deposition of 7 and 10 kg N ha⁻¹ per year were compared. Number of species was lower in the regions with the highest deposition. Literature data for plants on N concentration, nitrate reductase activity (NRA), growth rates, morphology and height were calculated on a site basis. They varied to different extent between the regions. The N concentration was 7–24% higher in the regions with the highest N deposition. We argue that the effect-related critical load based on our results should be set to a N deposition of 7–10 kg N ha⁻¹ per year. Critical loads for a subdivision of deciduous forests would give lower critical loads for the most acid soils compared to less acid soil.     

Importance of weed control, fertilization, irrigation, and genetics in slash and loblolly pine early growth on poorly drained spodosols

Research reported here has evaluated interactions of factors limiting the biological potential of slash pine (Pinus elliottii var. elliottii Engelm.) and loblolly pine (P. Taeda L.). Water and nutrients were manipulated by irrigation, weed control, and intensive fertilization. Genetic factors were incorporated as (1) improved loblolly seedlings from a commercial clonal seed orchard and (2) slash seedlings from four fast-growing, open-pollinated seed-orchard clones. Three replications of a 2 × 2 × 2 factorial experiment were established on a Typic Haplaquod soil in northern Florida using four cultural treatments: no treatment (F0H0); weed control (F0H1); fertilizer (F1F0); and combination of weed control and fertilizer (F1H1) for each species. During the first year there were large, sustained responses to all cultural practices. Loblolly and slash pine showed similar growth responses and had a lengthened growing season (60–100 days). Slash pine growth was superior to that of loblolly. Growth responses observed during the first year continued. After 4 years, irrigation had no measurable effect on tree growth. Both pine species responded equally to F1H0 and F0H1, but slash retained a slight edge. Extra nutrients]available via fertilizer, reduced competition, or additional rooting space, increased slash-pine volume index from an average of 2.0 m³ ha⁻¹ (F0H0) to 11.90 and 13.59 m³ ha⁻¹ (F0H1 and F1H0, respectively). The F1H1 treatment produced the largest slash-pine response, diameter at breast height (D_bh) averaged 9.84 cm, height reached 5.02 m, and volume index rose to about 22.94 m³ ha⁻¹. With teh F1H1 treatment, average loblolly-pine height and volume exceeded that of the average slash pine. At age 4, the growth response due to increased nutrient supply was 13–29 times that of the best genetic response within F0H0 trees of the four slash-pine progenies. The best-growing slash-pine family (6–56) exceeded loblolly-pine growth under all four treatments and approximated the vigor shown by slash pine in the southern hemisphere.