Dissolved Al reduces Mg uptake in Norway spruce forest: Results from a long-term field manipulation experiment in Norway

Dissolved aluminium (Al) in soils, mobilized by acid deposition, is considered a threat to forest health through hampering root growth and nutrient uptake. Since the end of the 1980s dissolved Al in forest soil water plays a key role in the assessment of critical loads of acid deposition. So far, most evidence for toxicity of dissolved Al in forest soil water is based on nutrient solution studies and pot experiments. Here, we present results from one of the few in situ ecosystem-scale forest manipulation experiments to study the effect of Al on mature forest trees. A plotwise addition of dilute AlCl₃ was conducted during seven years in an even-aged spruce forest (Picea abies) in an area in Norway with low acid deposition. Soil solution concentrations of Al were increased to potentially toxic levels (up to 500 μmol L⁻¹) and base cation (Ca + Mg + K) to inorganic Al ratios in the soil solution in the root zone were mostly below 1 in the Al-addition treatments. In the control treatment (only water addition) Al concentrations did not exceed 15 μmol L⁻¹ and base cation to inorganic Al ratios were above 1. The toxic effects of Al on fine root growth and plant growth found in hydroponic studies and pot trials are not confirmed by this field manipulation. However, magnesium (Mg) contents in needles decreased significantly and persistently in plots with elevated Al concentrations, whereas the needle Ca content did not respond. The depletion of the Mg content in needles is suggested to be due to antagonistic effects of high Al concentrations at the root surface, consistent with observed reductions in Mg to Al ratio of inner bark. This study clearly supports a role for Al in critical load functions for forests as dissolved Al causes a decrease in uptake of Mg. However, other signs of reduced forest vitality were not observed. Soil base cation status may need to be included in risk evaluations of forest health under acid deposition.     

Nutrient concentrations in roots, leaves and wood of seedling and mature sugar maple and American beech at two contrasting sites

Differences in sensitivity to soil conditions across tree species and developmental stage are important to predicting forest response to environmental change. This study was conducted to compare elemental concentrations in leaves, stems, and roots of (1) sugar maple (Acer saccharum Marsh.) seedlings vs. mature trees and (2) mature sugar maple vs. mature American beech (Fagus grandifolia Ehrh.) in two sites that differ in soil base saturation and pH. Both sites are located in Huntington Forest, NY, USA; one site (hereafter ‘H’) has higher soil pH and Ca, Mg, and Mn concentrations than the other site (hereafter ‘L’). Sugar maple growth at H (14.8 cm² year⁻¹ per tree) was much greater than at L (8.6 cm² year⁻¹ per tree), but the growth of beech was not different between the two sites. Leaves, roots, and stem wood of mature beech trees and sugar maple seedlings and mature trees were sampled for nutrient analysis. Foliar Ca, K, and Al concentrations were positively correlated with soil elements, but Mn concentrations were negatively correlated. Sugar maple differed more than beech between sites in foliar K and Mn concentrations. Root Mg and P concentrations reflected soil chemistry differences, in contrast to foliar concentrations of Mg and P, which were indistinguishable between the sites. In sugar maple, seedlings differed more than in mature trees in nutrient concentrations in roots, especially for Mg and Mn. Although beech was not as responsive to nutrient availability as sugar maple in foliar and root nutrient concentrations, Ca and Mg concentrations in beech wood were higher in H (52% higher for Ca and 68% higher for Mg), while sugar maple did not differ between sites. Sugar maple regeneration failure on acidic soils in the same region is consistent with our finding that sugar maple seedlings were very sensitive to nutrient availability. This sensitivity could ultimately contribute to the replacement of sugar maple by American beech in regions of low pH and base cations if base cation leaching by anthropogenic deposition and tree harvesting continues.     

Aluminium concentration versus the base cation to aluminium ratio as predictors for aluminium toxicity in Pinus sylvestris and Picea abies seedlings

Aluminium (Al) toxicity is considered an important factor in forest deterioration caused by soil acidification. A ratio of base cations (BC) to Al in the soil solution lower than 1 is widely used as an indicator for potentially adverse effects on tree health. In our view, the validity of the assumptions underlying the use of the BC:Al ratio as an indicator for Al toxicity in trees has never been evaluated properly.Here, we evaluate the importance of the base cations Ca and Mg in counteracting Al toxicity. Pinus sylvestris and Picea abies seedlings were grown on nutrient solution with a range of Al (0–0.25–0.5–1–2 mM) and base cation (0.25–0.5–2 mM) concentrations, giving BC:Al ratios of 1 at different levels of Al. Increasing concentrations of Al in solution caused growth reductions, which could not be counteracted by increasing concentrations of BC in solution with P. sylvestris and only partly counteracted with P. abies. Increased concentrations of Al in solution decreased the concentrations in shoot and root of both Ca and Mg, while increased concentrations of BC in solution increased tissue concentrations of BC. Growth reductions were, however, not a result of BC deficiencies, as growth reduction already occurred in tree seedlings that maintained adequate concentrations of Ca and Mg.All growth and uptake variables measured showed a higher or equal correlation with the absolute concentrations of Al or Al+BC in solution than with the BC:Al ratio. We conclude that Al toxicity is determined solely by the concentration of Al in solution. Shoot growth decreased significantly as dissolved Al increased at a constant BC:Al ratio of 1. In P. abies, but not in P. sylvestris, dissolved BC can positively affect uptake of BC and growth, which might partly alleviate the toxic effects of Al. Our results show that the mechanistic explanation for the effect of the BC:Al ratio is insufficient to describe Al toxicity. Care should be taken when using models based on the BC:Al ratio to predict the effect of Al on tree growth.     

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

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

Sugar maple and yellow birch regeneration in response to canopy opening,liming and vegetation control in a temperate deciduous forest of Quebec

We examined how the density, growth and survival of sugar maple (Acer saccharum Marsh.) and yellow birch (Betula alleghaniensis Britton) regeneration are influenced by gap size, soil nutrient availability and understory vegetation. We used a factorial combination of (1) three gap sizes (small: <100 m²; medium: 100–300 m²; large: ∼1000 m²); (2) presence/absence of liming (92% CaCO₃ at 500 kg ha⁻¹, 1st year post-harvest); and (3) presence/absence of vegetation control (weeding twice a year; 1st to 3rd year post-harvest). We monitored height increment and survival of 1500 seedlings and saplings of both species from the 3rd to the 6th year post-harvest, and assessed density 6 years post-harvest. Both species exhibited a complex set of density, growth and survival responses across the combination of treatments. Compared to sugar maple, yellow birch had an overall lower density, greater growth, and similar survival rate; the two species attained maximum values in different gap size for density, and similar gap size for growth and survival. Liming had very little or no effect on the species. The growth of yellow birch was slightly but significantly greater when understory vegetation was controlled, particularly in medium and large gaps. These results suggest that a variety of canopy gap sizes can provide the right combination of understory conditions for regenerating these two functionally different tree species.     

An ecosystem approach to biodiversity effects: Carbon pools in a tropical tree plantation

This paper presents a synthesis of experiments conducted in a tropical tree plantation established in 2001 and consisting of 22 plots of 45 m × 45 m with either one, three or six native tree species. We examined the changes in carbon (C) pools (trees, herbaceous vegetation, litter, coarse woody debris (CWD), and mineral topsoil at 0-10 cm depth) and fluxes (decomposition of CWD and litter, as well as soil respiration) both through time and among diversity levels. Between 2001 and 2009 the aboveground C pools increased, driven by trees. Across diversity levels, the mean observed aboveground C pool was 7.9 ± 2.5 Mg ha⁻¹ in 2006 and 20.4 ± 7.4 Mg ha⁻¹ in 2009, a 158% increase. There was no significant diversity effect on the observed aboveground C pool, but we found a significant decrease in the topsoil C pool, with a mean value of 34.5 ± 2.4 Mg ha⁻¹ in 2001 and of 25.7 ± 5.7 Mg ha⁻¹ in 2009 (F_1,36 = 52.12, p < 0.001). Assuming that the biomass C pool in 2001 was negligible (<1 Mg ha⁻¹), then the plantation gained in C, on average, ∼20 and lost ∼9 Mg ha⁻¹ in biomass and soil respectively, for an overall gain of ∼11 Mg ha⁻¹ over 8 years. Across the entire data set, we uncovered significant effects of diversity on CWD decomposition (diversity: F_2,393 = 15.93, p < 0.001) and soil respiration (monocultures vs mixtures: t = 15.35, df = 11, p < 0.05) and a marginally significant time × diversity interaction on the loss of total C from the mineral topsoil pool (see above). Monthly CWD decomposition was significantly faster in monocultures (35.0 ± 24.1%) compared with triplets (31.3 ± 21.0%) and six-species mixtures (31.9 ± 26.8%), while soil respiration was higher in monocultures than in mixtures (t = 15.35, df = 11, p < 0.001). Path analyses showed that, as diversity increases, the links among the C pools and fluxes strengthen significantly. Our results demonstrate that tree diversity influences the processes governing the changes in C pools and fluxes following establishment of a tree plantation on a former pasture. We conclude that the choice of tree mixtures for afforestation in the tropics can have a marked influence on C pools and dynamics.     

Fine root production and turnover in forest ecosystems in relation to stand and environmental characteristics

The production and turnover of fine roots (diameter ?2 mm) contributes significantly to carbon cycling in forest ecosystems. We compiled an up-to-date global database covering 186 stands from the literature and estimated fine root production (FRP) and fine root turnover (FRT) for boreal, temperate and tropical forests in order to study the relationships between FRP or FRT and environmental and stand variables. FRP for all plants (trees + understorey) was 311 ± 259 (n = 39), 428 ± 375 (n = 71) and 596 ± 478 g m⁻² a⁻¹ (n = 32) in the boreal, temperate and tropical forests, respectively, and the corresponding annual FRT rates were 0.77 ± 0.70, 1.21 ± 1.04 and 1.44 ± 0.76, respectively. When the FRP and FRT of trees were estimated separately for boreal and temperate forests the differences between the two biomes were insignificant. The mean FRP of trees for the two biomes combined was 306 ± 240 g m⁻² a⁻¹ (n = 86) and the annual FRT was 1.31 ± 1.43. Fine root biomass (FRB) was the most significant factor explaining the variation in FRP, and more so at the tree level than at the stand level, explaining 53% of the variation in FRP for trees at the tree level. The corresponding proportions at the stand level were 21% for all plants and 12% for trees. Latitude, mean annual temperature and annual precipitation each explained <20% of the variation in FRP or FRT. Fine root production and FRT estimates are highly dependent on the species included in the sampling, the sampling depth and the methods used for estimating FRP or calculating FRT. The results indicate that the variation in FRP on a global scale can be explained to a higher degree if we focus on tree roots separately from the roots of the understorey vegetation and on FRP at the tree level instead of FRP at the stand level or on FRT.     

The impact of atmospheric deposition and climate on forest growth in European monitoring plots: An individual tree growth model

In the climate change discussion, the possibility of carbon sequestration of forests plays an important role. Therefore, research on the effects of environmental changes on net primary productivity is interesting. In this study we investigated the influence of changing temperature, precipitation and deposition of sulphur and nitrogen compounds on forest growth. The database consisted of 654 plots of the European intensive monitoring program (Level II plots) with 5-year growth data for the period 1994–1999. Among these 654 plots only 382 plots in 18 European countries met the requirements necessary to be used in our analysis. Our analysis was done for common beech (Fagus sylvatica), oak (Quercus petraea and Q. robur), Scots pine (Pinus sylvestris) and Norway spruce (Picea abies). We developed an individual tree growth model with measured basal area increment of each individual tree as responding growth factor and tree size (diameter at breast height), tree competition (basal area of larger trees and stand density index), site factors (soil C/N ratio, temperature), and environmental factors (temperature change compared to long-term average, nitrogen and sulphur deposition) as influencing parameters. Using a mixed model approach, all models for the tree species show a high goodness of fit with Pseudo-R² between 0.33 and 0.44. Diameter at breast height and basal area of larger trees were highly influential variables in all models. Increasing temperature shows a positive effect on growth for all species except Norway spruce. Nitrogen deposition shows a positive impact on growth for all four species. This influence was significant with p < 0.05 for all species except common beech. For beech the effect was nearly significant (p = 0.077). An increase of 1 kg N ha⁻¹ yr⁻¹ corresponds to an increase in basal area increment between 1.20% and 1.49% depending on species. Considering an average total carbon uptake for European forests near 1730 kg per hectare and year, this implies an estimated sequestration of approximately 21–26 kg carbon per kg nitrogen deposition.     

Acid deposition effects on forest composition and growth on the Monongahela National Forest,West Virginia

The northern and central Appalachian forests are subject to high levels of atmospheric acid deposition (AD), which has been shown in some forests to negatively impact forest growth as well as predispose the forest system to damage from secondary stresses. The purpose of this study was to evaluate the possible contribution of AD to changes in composition and productivity of the Monongahela National Forest, and to evaluate soil-based indicators of acidification that might be useful for detecting AD-related forest changes. Soils adjacent to 30 Forest Inventory and Analysis (FIA) sites were sampled and analyzed for a suite of acidity indicators. These indicators were correlated with the periodic mean annual volume increment (PMAVI) of the forest stands on FIA plots for the 10-yr period 1989–2000. PMAVI ranged from −9.5 to 11.8 m³ ha⁻¹ yr⁻¹, with lower-than-expected growth (<3 m³ ha⁻¹ yr⁻¹) on two-thirds of the sites. In the surface horizon, effective base saturation, Ca²⁺ concentration, base saturation, K⁺ concentration, Ca/Al molar ratio, and Mg/Al molar ratio, were positively correlated with PMAVI and Fe concentration was negatively correlated with PMAVI (p ≤ 0.1). In the subsurface horizon pH_(w) and effective base saturation were positively correlated and Al³⁻ concentration and K⁺ concentration were negatively correlated with PMAVI. We hypothesized that NO₃-N/NH₄-N ratio would also be correlated with PMAVI, but it was not. Correlations between soil chemical indicators and PMAVI suggest that AD may contribute, in part, to the lower-than-expected forest growth on the Monongahela National Forest.     

Robust relationships for simple plantation growth models based on sparse data

Three equations predicting height H = β₁(t − 0.5)^0.5, diameter D = β₂(H − 1.3)/ln N, and mortality dN/N = −2(G/G_max)³dD/D from plantation age (t), stocking (N) and basal area (G) can be calibrated with few data (even a single observation) for plantations in which re-measured data and growth models are unavailable. Despite having only three parameters to be estimated, these equations extrapolate reliably and allow objective forecasts of future plantation growth performance that may serve as useful first approximations until more precise growth models can be developed.     

Above- and belowground biomass in a Brazilian Cerrado

Cerrado is a biome that occupies about 25% of the Brazilian territory and is characterized by a gradient of grassland to savanna and forest formations and by high species richness. It has been severely affected by degradation and deforestation and has been heavily fragmented over the past 4-5 decades. Despite the recognized overall ecological importance of the Cerrado, there are only few studies focusing on the quantification of biomass in this biome. We conducted such a case study in the South-East of Brazil in a cerrado sensu stricto (cerrado s.s.) with the goal to produce estimates of above- and belowground biomass and to develop allometric equations. A number of 120 trees from 18 species were destructively sampled and partitioned into the components: leaves, branches and bole. Five models with DBH (D), height (H), D²H and wood density (WD) as independent variables were tested for the development of allometric models for individual tree aboveground biomass (leaves + branches + bole). One model based on basal area (BA) as a stand parameter was also tested as an alternative approach for predicting aboveground biomass in the stand level. Belowground biomass was estimated by subsampling on 10 sample plots. Mean aboveground tree biomass (bole, branches and leaves) was estimated to be 62,965.5 kg ha⁻¹(SE = 14.6%) and belowground biomass accounted for 37,501.8 kg ha⁻¹ (SE = 23%). The best-fit equation for the estimation of individual tree aboveground biomass include DBH and wood density as explanatory variables (R² = 0.898; SEE = 0.371) and is applicable for the diameter range of this study (5.0-27.6 cm) and in environments with similar conditions of the cerrado s.s. sampled. In the stand level, the model tested presented a higher goodness of fit than the single tree models (R² = 0.934; SEE = 0.224). Our estimates of aboveground biomass are higher than reported by other studies developed in the same physiognomy, but the estimates of belowground biomass are within the range of values reported in other studies from sites in cerrado s.s. Both biomass estimates, however, exhibit relatively large standard errors. The root-to-shoot ratio of the sample trees is in the magnitude of reported values for savanna ecosystems, but smaller than estimated from other studies in the cerrado s.s.     

Carbon concentration variability of 10 Chinese temperate tree species

A mass-based carbon (C) concentration ([C]) of 50% in dry wood is widely accepted as a constant factor for conversion of biomass to C stock. However, the [C] varies with tree species, and few data on [C] are available for the Chinese temperate tree species. In this study, we examined inter- and intra-specific variations of [C] in biomass tissues for 10 co-occurring temperate tree species in northeastern China. The species were Korean pine (Pinus koraiensis Sieb. et Zucc.), Dahurian larch (Larix gmelinii Rupr.), Mongolian oak (Quercus mongolica Fisch.), white birch (Betula platyphylla Suk.), Amur cork-tree (Phellodendron amurense Rupr.), Manchurian walnut (Juglans mandshurica Maxim.), Manchurian ash (Fraxinus mandshurica Rupr.), aspen (Populous davidiana Dode), Mono maple (Acer mono Maxim.), and Amur linden (Tilia amurensis Rupr.). The mean tissue [C] across the species varied from 47.1% in fine root to 51.4% in foliage. The mean stem [C] of the 10 species was 49.9 ± 1.3% (mean ± SE). The weighted mean C concentration (WMCC) for the species ranked as: Amur cork-tree (55.1%) > Amur linden (53.9%) > Korean pine (53.2%) > Manchurian ash (52.9%) > Manchurian walnut (52.4%) > Mongolian oak (47.6%) > Dahurian larch (46.9%) > Mono maple (46.4%) > white birch (46.1%) > aspen (43.7%). The WMCC of the dominant trees was negatively correlated to mean annual increment of biomass (MAI), suggesting that planting fast-growing tree species for C sequestration in afforestation and reforestation practices sacrifice some C gain from increasing MAI due to decreasing [C]. Failing to account for the inter- and intra-specific variations in [C] will introduce a relative error of −6.7% to +7.2% in estimates of biomass C stock from inventory data, of which >93% is attributed to ignoring the inter-specific variation in [C].     

Recovery of carbon and nutrient pools in a northern forested wetland 11 years after harvesting and site preparation

We measured the change in above- and below-ground carbon and nutrient pools 11 years after the harvesting and site preparation of a histic-mineral soil wetland forest in the Upper Peninsula of Michigan. The original stand of black spruce (Picea mariana), jack pine (Pinus banksiana) and tamarack (Larix laricina) was whole-tree harvested, and three post-harvest treatments (disk trenching, bedding, and none) were randomly assigned to three Latin square blocks (n = 9). Nine control plots were also established in an adjoining uncut stand. Carbon and nutrients were measured in three strata of above-ground vegetation, woody debris, roots, forest floor, and mineral soil to a depth of 1.5 m. Eleven years following harvesting, soil C, N, Ca, Mg, and K pools were similar among the three site preparation treatments and the uncut stand. However, there were differences in ecosystem-level nutrient pools because of differences in live biomass. Coarse roots comprised approximately 30% of the tree biomass C in the regenerated stands and 18% in the uncut stand. Nutrient sequestration, in the vegetation since harvesting yielded an average net ecosystem gain of 332 kg N ha⁻¹, 110 kg Ca ha⁻¹, 18 kg Mg ha⁻¹, and 65 kg K ha⁻¹. The likely source for the cations and N is uptake from shallow groundwater, but N additions could also come from non-symbiotic N-fixation and N deposition. These are the only reported findings on long-term effects of harvesting and site preparation on a histic-mineral soil wetland and the results illustrate the importance of understanding the ecohydrology and nutrient dynamics of the wetland forest. This wetland type appears less sensitive to disturbance than upland sites, and is capable of sustained productivity under these silvicultural treatments.     

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

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

Forest thinning and subsequent bark beetle-caused mortality in Northeastern California

The Warner Mountains of northeastern California on the Modoc National Forest experienced a high incidence of tree mortality (2001–2007) that was associated with drought and bark beetle (Coleoptera: Curculionidae, Scolytinae) attack. Various silvicultural thinning treatments were implemented prior to this period of tree mortality to reduce stand density and increase residual tree growth and vigor. Our study: (1) compared bark beetle-caused conifer mortality in forested areas thinned from 1985 to 1998 to similar, non-thinned areas and (2) identified site, stand and individual tree characteristics associated with conifer mortality. We sampled ponderosa pine (Pinus ponderosa var ponderosa Dougl. ex Laws.) and Jeffrey pine (Pinus jeffreyi Grev. and Balf.) trees in pre-commercially thinned and non-thinned plantations and ponderosa pine and white fir (Abies concolor var lowiana Gordon) in mixed conifer forests that were commercially thinned, salvage-thinned, and non-thinned. Clusters of five plots (1/50th ha) and four transects (20.1 × 100.6 m) were sampled to estimate stand, site and tree mortality characteristics. A total of 20 pre-commercially thinned and 13 non-thinned plantation plot clusters as well as 20 commercially thinned, 20 salvage-thinned and 20 non-thinned mixed conifer plot clusters were established. Plantation and mixed conifer data were analyzed separately. In ponderosa pine plantations, mountain pine beetle (Dendroctonus ponderosae Hopkins) (MPB) caused greater density of mortality (trees ha⁻¹ killed) in non-thinned (median 16.1 trees ha⁻¹) compared to the pre-commercially thinned (1.2 trees ha⁻¹) stands. Percent mortality (trees ha⁻¹ killed/trees ha⁻¹ host available) was less in the pre-commercially thinned (median 0.5%) compared to the non-thinned (5.0%) plantation stands. In mixed conifer areas, fir engraver beetles (Scolytus ventralis LeConte) (FEN) caused greater density of white fir mortality in non-thinned (least square mean 44.5 trees ha⁻¹) compared to the commercially thinned (23.8 trees ha⁻¹) and salvage-thinned stands (16.4 trees ha⁻¹). Percent mortality did not differ between commercially thinned (least square mean 12.6%), salvage-thinned (11.0%), and non-thinned (13.1%) mixed conifer stands. Thus, FEN-caused mortality occurred in direct proportion to the density of available white fir. In plantations, density of MPB-caused mortality was associated with treatment and tree density of all species. In mixed conifer areas, density of FEN-caused mortality had a positive association with white fir density and a curvilinear association with elevation.     

Effects of winter selective tree harvest on soil microclimate and surface CO2 flux of a northern hardwood forest

Soil surface CO₂ flux (S_flux) is the second largest terrestrial ecosystem carbon flux, and may be affected by forest harvest. The effects of clearcutting on S_flux have been studied, but little is known about the effect of alternative harvesting methods such as selective tree harvest on S_flux. We measured S_flux before and after (i) the creation of forest canopy gaps (simulating group tree selection harvests) and (ii) mechanized winter harvest but no tree removal (simulating ground disturbance associated with logging). The experiment was carried out in a sugar maple dominated forest in the Flambeau River State Forest, Wisconsin. Pre-treatment measurements of soil moisture, temperature and S_flux were measured throughout the growing season of 2006. In January–February 2007, a harvester created the canopy gaps (200–380 m²). The mechanization treatment consisted of the harvester traveling through the plots for a similar amount of time as the gap plots, but no trees were cut. Soil moisture and temperature and S_flux were measured throughout the growing season for 1 year prior to harvest and for 2 years after harvest. Soil moisture and temperature were significantly greater in the gap than mechanized and control treatments. Instantaneous S_flux was positively correlated to soil moisture and soil temperature at 2 and 10 cm, but temperature at 10 cm was the single best predictor. Annual S_flux was not significantly different among treatments prior to winter 2007 harvest, and was not significantly different among treatments after harvest. Annual (+1 std. err.) S_flux averaged 967 + 72, 1011 + 72, and 1012 + 72 g C m⁻² year⁻¹ in the control, mechanized and gap treatments, respectively, for the 2-year post-treatment period. The results from this study suggest selective group tree harvest significantly increases soil moisture and temperature but does not significantly influence S_flux.     

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

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

Measuring heights to crown base and crown median with LiDAR in a mature,even-aged loblolly pine stand

This study evaluated the possibility of measuring the height to the base of the live crown and the height to the median of canopy elements with airborne scanning LiDAR (Light Detection And Ranging) in a simple, even-aged stand of loblolly pine. The first step in determining these heights was fitting truncated Weibull functions to the vertical distribution of elevations where discrete laser pulses were reflected from the dominant canopy strata. The height to the canopy median was defined as height at the median of the distribution. The height to the base of the live crown was defined as the height where the upper tail the distribution asymptotes to zero returns. Ground-based and LiDAR-based estimates of the canopy median differed by 0.3 m and were not significantly different (P = 0.23). Ground- and LiDAR-based estimates of the base of the live crown differed by 0.6 m and were significantly different (P = 0.03). LiDAR-based estimates of the canopy median exhibited positive bias over most of the range of field-measured values. Analyses of the LiDAR data resulted in overestimating the height to the canopy base over most of the range in field-measured values; however, the difference between ground and LiDAR-based estimates were negatively correlated with ground-based measurements. Average tree diameter was calculated with LiDAR-generated heights to the canopy median and to the base of the live crown. The overall average diameter was not statistically different from the overall quadratic mean diameter measured on the ground, demonstrating the possible utility of these canopy variables to forest managers working with simple stands such as plantations.     

Spatial variability in stand structure and density-dependent mortality in newly established post-fire stands of a California closed-cone pine forest

Fire is an important process in California closed-cone pine forests; however spatial variability in post-fire stand dynamics of these forests is poorly understood. The 1995 Vision Fire in Point Reyes National Seashore burned over 5000 ha, initiating vigorous Pinus muricata (bishop pine) regeneration in areas that were forested prior to the fire but also serving as a catalyst for forest expansion into other locales. We examined the post-fire stand structure of P. muricata forest 14 years after fire in newly established stands where the forest has expanded across the burn landscape to determine the important factors driving variability in density, basal area, tree size, and mortality. Additionally, we estimated the self-thinning line at this point in stand development and compared the size-density relationship in this forest to the theorized (−1.605) log-log slope of Reineke’s Rule, which relates maximum stand density to average tree size. Following the fire, post-fire P. muricata density in the expanded forest ranged from 500 to 8900 live stems ha⁻¹ (median density = 1800 ha⁻¹). Post-fire tree density and basal area declined with increasing distance to individual pre-fire trees, but showed little variation with other environmental covariates. Self-thinning (density-dependent mortality) was observed in nearly all stands with post-fire density >1800 stems ha⁻¹, and post-fire P. muricata stands conformed to the size-density relationship predicted by Reineke’s Rule. This study demonstrates broad spatial variability in forest development following stand-replacing fires in California closed-cone pine forests, and highlights the importance of isolated pre-fire trees as drivers of stand establishment and development in serotinous conifers.     

Organic residue mass at planting is an excellent predictor of tree growth in Eucalyptus plantations established on a sandy tropical soil

Tropical plantation forests are meeting an increasing proportion of global wood demand and comprehensive studies assessing the impact of silvicultural practices on tree and soil functioning are required to achieve sustainable yields. The objectives of our study were: (1) to quantify the effects of contrasting organic residue (OR) retention methods on tree growth and soil nutrient pools over a full Eucalyptus rotation and (2) to assess the potential of soil analyses to predict yields of fast-growing plantations established on tropical sandy soils. An experiment was set up in the Congo at the harvesting of the first rotation after afforestation of a native herbaceous savanna. Six treatments were set up in 0.26 ha plots and replicated in 4 blocks, with OR mass at planting ranging from 0 to 46.5 Mg ha⁻¹. Tree growth over the whole rotation was highly dependent on OR management at planting. Over-bark trunk volume 7 years after planting ranged from 96 m³ ha⁻¹ in the treatment with forest floor and harvest residue removal at planting to 164 m³ ha⁻¹ in the treatment with the largest amount of OR. A comparison of nutrient stocks within the ecosystem at planting and at the end of the rotation suggested that nutrient contents in OR were largely involved in the different response observed between treatments. OR management treatments did not significantly modify most of the nutrient concentrations in the upper layers of the mineral soil. Conventional soil analyses performed before planting and at ages 1 and 3 years were unable to detect differences between treatments despite large differences in tree growth. In contrast, linear regressions between stand aboveground biomass at harvesting and OR mass at planting (independent variable) showed that OR mass was an excellent predictor of stand yield (R² = 0.99). A large share of soil fertility comes from organic material above the mineral soil in highly weathered sandy soils and OR mass at planting might be used in conjunction with soil analyses to assess the potential of these soils to support forest plantations.