Diversity of northern plantations peaks at intermediate management intensity

Tree diversity is an important component of biodiversity. Management intensification is hypothesized to affect tree diversity. However, evidence to support the relationship between management intensity and tree diversity in northern forests is lacking. This study examined the effects of fertilization, site preparation, and brush control on tree species diversity, shade tolerance diversity and size diversity of jack pine (Pinus banksiana Lamb.), black spruce (Picea mariana [Mill.] B.S.P.), white pine (Pinus strobus L.) and white spruce (Picea glauca [Moench] Voss) plantations, 15 years after planting in Ontario, Canada. Species diversity and shade tolerance diversity were highly correlated, so were diameter size diversity and height size diversity. Fertilization did not affect the tree diversity indices of any plantations. Species diversity and shade tolerance diversity was interactively influenced by site preparation and brush control in the black spruce, white pine, and white spruce plantations, showing that the highest diversity occurred on sites with intensive site preparation without brush control, whereas on sites with brush control, diversity was higher with least intensity of site preparation. However, in the jack pine plantation, neither species diversity nor shade tolerance diversity differed with management intensification, and is attributed to the fast capture of site resources by the planted crop trees of jack pine which minimized establishment of non-crop species. Tree size diversity increased with site preparation intensity in the jack pine and black spruce plantations, while it decreased with brush control in the white pine and white spruce plantations. We concluded that (1) the effects of management intensification on diversity of northern plantations differ with growth habit of planted crop tree species and (2) species diversity and tree size diversity tend to be highest at intermediate levels of silvicultural intensification during the stand establishment phase, supporting the intermediate disturbance hypothesis.     

Predicting canopy macrolichen diversity and abundance within old-growth inland temperate rainforests

Windward slopes of the inland mountain ranges in British Columbia support a unique inland temperate rainforest (ITR) ecosystem. Increasing fragmentation and the loss of old ITR stands have highlighted the need for determining conservation biology priorities among remaining old forest stands. We have addressed this issue by surveying foliose macrolichens within 53 old ITR stands in British Columbia's 135,000 ha very wet-cool interior-cedar hemlock (ICHvk2) biogeoclimatic subzone in the upper Fraser River watershed. Study plots were stratified by leading tree species and by “wet” versus “dry” relative soil moisture conditions. Other plot variables included: temperature, precipitation, incident solar loading, and canopy openness. Ordination plots showed a distinct assemblage of foliose cyanolichens, including Lobaria pulmonaria, Lobaria retigera, Sticta fuliginosa, Nephroma isidiosum, Nephroma occultum, and Pseudocyphellaria anomala, whose abundance was correlated with increasing relative soil moisture, temperature, canopy openness, precipitation, and basal area of spruce. Logistic regression models similarly identified relative soil moisture and temperature in all parsimonious models. Leading tree species, in combination with “wet” relative soil moisture and/or temperature, were important factors explaining the presence or absence of five (Cavernularia hultenii, L. retigera, N. occultum, Platismatia norvegica, and Sticta oroborealis) of the eight modeled old-growth associate lichen species. This combination of conditions favouring the development of canopy lichen communities in old forests was best expressed in low elevation water receiving sites. We hypothesize that groundwater availability in these sites promotes species richness and abundance of canopy lichens by creating more favourable conditions for growth, and by reducing fire return intervals which allows for the accumulation of rare species over time. Historically, forests in these wet “toe-slope” positions were disproportionately targeted for logging. Their conservation should now be a high priority, given their disproportionate significance to maintaining canopy lichen diversity in the present-day landscapes.     

Changes in the canopies of Pinus sylvestris and Picea abies under alkaline dust impact in the industrial region of Northeast Estonia

Studies were carried out in 1999, 2005 and 2007 in the area of Kunda cement plant in Northeast Estonia on sample plots 3 km W and 2.5 and 5 km E of Kunda. As control stands, two plots for pine and spruce were established in Lahemaa National Park (34-38 km W of Kunda). The selected pine and spruce stands were 75-85-year-old Myrtillus site type, of 0.7-0.8 density and II quality class, with moderately dense or sparse understorey. The values concerning needle density and number of needle scars were higher for shoots formed in the period of higher pollution than for the shoots grown under a considerably lower pollution load. Although the cement dust pollution has notably decreased from year to year, the number of needle pairs per 1 cm of the shoot was 1.8-2.1 times greater in the shoots formed in 1998 than in those formed in 2003, whereas the changes were statistically reliable. Possibly the low temperatures at the time of shoot and needle formation affected the density of needles on all sample plots, and thus the number of needles on shoots formed in 2003 was many times smaller. After the significant fall in the pollution load since 1996 the length growth of needles intensified around Kunda cement plant, at the same time no changes occurred in the length growth of needles in the control area. As compared to the data from 1998, the length growth of pine needles had improved, especially 2.5 and 5 km E from the cement plant, needles being respectively 1.5 and 1.1 cm longer than 6 years ago. The stimulation of the growth of pine and spruce needles 2.5 and 5 km E of the cement plant may be a sign of a positive effect of reduced doses of cement dust in soil. The greater length of pine and spruce needles is the reason for the larger biomass of the needles.     

Spatial impacts of soil disturbance and residual overstory on density and growth of regenerating aspen

We examined spatial aspects of harvesting impacts on aspen regeneration at 25 sites in northern Minnesota. These sites had been clearcut or partially harvested 4–11 years ago. At each site, residual overstory, which was composed of trees other than aspen, soil disturbance, and tree regeneration were determined along transects leading away from skid trails into the neighboring stand. We characterized spatial extent of soil disturbance as soil strength using an Eijkelkamp soil cone penetrometer. Soil disturbance dropped off very quickly at the edge of skid trails, suggesting that the impact of harvesting traffic on areas adjacent to skid trails is minor. On skid trails, disturbance levels were higher on sites harvested in summer than on sites harvested in winter. Even after adjustment for differences in soil disturbance, stands harvested in winter had higher regeneration densities and greater aspen height growth than stands harvested in summer, suggesting that aspen regeneration was more sensitive to a given level of soil disturbance on summer-harvested sites versus on winter-harvested sites. Soil disturbance and residual overstory interactively reduced aspen regeneration densities and height growth, indicating that avoidance of soil disturbance is even more critical in partially harvested stands. Predictions based in the spatial patterns of impact found in this study indicated that harvesting conditions may have a great impact in future productivity of a site.     

Differences in soil properties in adjacent stands of Scots pine, Norway spruce and silver birch in SW Sweden

Soil properties were compared in adjacent 50-year-old Norway spruce, Scots pine and silver birch stands growing on similar soils in south-west Sweden. The effects of tree species were most apparent in the humus layer and decreased with soil depth. At 20-30 cm depth in the mineral soil, species differences in soil properties were small and mostly not significant. Soil C, N, K, Ca, Mg, and Na content, pH, base saturation and fine root biomass all significantly differed between humus layers of different species. Since the climate, parent material, land use history and soil type were similar, the differences can be ascribed to tree species. Spruce stands had the largest amounts of carbon stored down to 30 cm depth in mineral soil (7.3 kg C m⁻²), whereas birch stands, with the lowest production, smallest amount of litterfall and lowest C:N ratio in litter and humus, had the smallest carbon pool (4.1 kg C m⁻²), with pine intermediate (4.9 kg C m⁻²). Similarly, soil nitrogen pools amounted to 349, 269, and 240 g N m⁻² for spruce, pine, and birch stands, respectively. The humus layer in birch stands was thin and mixed with mineral soil, and soil pH was highest in the birch stands. Spruce had the thickest humus layer with the lowest pH.     

Growth and productivity of black spruce in even- and uneven-aged stands at the limit of the closed boreal forest

The increasing commercial interest and advancing exploitation of new remote territories of the boreal forest require deeper knowledge of the productivity of these ecosystems. Canadian boreal forests are commonly assumed to be evenly aged, but recent studies show that frequent small-scale disturbances can lead to uneven-aged class distributions. However, how age distribution affects tree growth and stand productivity at high latitudes remains an unanswered question. Dynamics of tree growth in even- and uneven-aged stands at the limit of the closed black spruce (Picea mariana) forest in Quebec (Canada) were assessed on 18 plots with ages ranging from 77 to 340 years. Height, diameter and age of all trees were measured. Stem analysis was performed on the 10 dominant trees of each plot by measuring tree-ring widths on discs collected each meter from the stem, and the growth dynamics in height, diameter and volume were estimated according to tree age. Although growth followed a sigmoid pattern with similar shapes and asymptotes in even- and uneven-aged stands, trees in the latter showed curves more flattened and with increases delayed in time. Growth rates in even-aged plots were at least twice those of uneven-aged plots. The vigorous growth rates occurred earlier in trees of even-aged plots with a culmination of the mean annual increment in height, diameter and volume estimated at 40–80 years, 90–110 years earlier than in uneven-aged plots. Stand volume ranged between 30 and 238 m³ ha⁻¹ with 75% of stands showing values lower than 120 m³ ha⁻¹ and higher volumes occurring at greater dominant heights and stand densities. Results demonstrated the different growth dynamics of black spruce in single- and multi-cohort stands and suggested the need for information on the stand structure when estimating the effective or potential growth performance for forest management of this species.     

Effects of non-native spruce plantations on small mammal communities in subarctic birch forests

Voles and shrews are key species in northern forest ecosystems. Thus, it is important to quantify to what extent new forestry practices such as planting of non-native tree species impact these small mammals. In northern Norway stands of coastal subarctic birch forests have increasingly been converted to non-native spruce stands during the last century. This leads to changes in the forest floor vegetation and soil conditions that can be expected to negatively impact the community of ground-dwelling small mammals. In this 10-year trapping study we contrasted seasonal small mammal population abundances in spruce plantations with four birch forest varieties. Six different small mammal species were trapped (in descending order of abundance; common shrew Sorex araneus, red vole Myodes rutilus, field vole Microtus agrestis, grey-sided vole M. rufocanus, pygmy shrew S. minutus and water shrew Neomys fodiens). None of the voles appeared to exhibit temporal dynamics resembling population cycles. The three most numerous species were clearly less abundant in the spruce plantations compared to the other forest types. Autumn abundances were most impacted by spruce plantations, indicating that growth rates in the reproductive season were more influenced than winter declines. Species associated with productive forest habitats (i.e. field vole and common shrew) were most impacted by tree species conversion. Still young spruce plantations inter-mixed with birch trees and the ecotone habitat, sustained small mammal abundances comparable to the native birch forests. This implies that managing spruce plantations to maintain a mix of different tree species and high spatial heterogeneity (i.e. more ecotones), will reduce the negative impacts on the small mammal community. On the contrary, if young spruce plantations, as they age become spruce monocultures covering larger parts of the landscapes than they do presently, the negative effects on small mammal communities may be larger than observed in the present study.     

Subsidy or stress? Tree structure and growth in wetland forests along a hydrological gradient in Southern Europe

In forested wetlands, hydrology exerts complex and sometimes compensatory influences on tree growth. This is particularly true in semi-arid ecosystems, where water can be both a limiting resource and a stressor. To better understand these relationships, we studied hydrologic and edaphic controls on the density, growth, tree architecture and overall productivity of forested wetlands dominated by the tree species Alnus glutinosa and Salix atrocinerea in Southern Europe. We sampled 49 plots set within 21 stands in the Atlantic coastal zone of the Iberian Peninsula, and quantified woody composition, size structure (diameter and height), and radial growth using dendrochronology. Plots were grouped into three saturation classes to compare tree growth characteristics (tree density, degree of sprouting, live basal area and productivity) across levels of saturation. We used Principal Component Analysis (PCA) to create integrated explanatory factors of hydrology, soil nutrient status and soil texture for use in linear mixed models to predict stand characteristics. Increased site saturation favoured a shift in species dominance from Alnus to Salix and resulted in a higher degree of multi-stemmed tree architecture (‘shrubbiness’), particularly for Alnus. Radial growth was negatively correlated with long-term soil saturation; however, annual productivity on a per-tree basis varied by species. Alnus growth and tree density were negatively correlated with waterlogging and fine-textured soils, possibly due to anaerobiosis in the rooting zone. In contrast, Salix growth was more influenced by nutrient limitation. Overall site productivity as measured by annual basal area increment decreased with prolonged saturation. In summary, soil saturation appears to act as a chronic stressor for tree species in this ecosystem. However, these species persist and maintain a dominant canopy position in the most permanently flooded patches through increased sprouting, albeit at a reduced rate of overall biomass accumulation relative to well-drained sites. The diversity in functional responses among wetland forest species has important implications for the conservation and management of these ecosystems. The sustainable management of these ecosystems is directly tied to their vulnerability to changing hydrological conditions. Non-equilibrium modifications to the hydrologic regime from land use and climate change, which are particularly severe in semi-arid regions, may further decrease productivity, integrity and resilience in these stress-adapted communities.     

Does mechanical site preparation affect trembling aspen density and growth 9–12 years after treatment?

Trembling aspen (Populus tremuloides Michx.) density and growth were assessed 9–12 years after stand establishment to determine whether mechanical site preparation (MSP) affects crop tree quality. Study sites were either treated with disc trenching or ripper ploughing and planted with white spruce (Picea glauca (Moench) Voss) seedlings immediately after harvest (treated) or were undisturbed since harvest (control). Stands were surveyed during the summer of 2002 with standard regeneration survey plots. Results show that aspen stem density and height were lower in MSP-treated areas relative to untreated areas. Diameter growth rates were unaffected by treatment, however the percentage of stem discolouration was higher in untreated control stands compared to site prepared areas. The results of this study indicate that there are no long-term benefits to carry out MSP for aspen promotion. However, as MSP does not appear to seriously harm the aspen crop, we suggest that this treatment can still be used on sites where aspen densities may be low without treatment (e.g., sites with extremely low soil temperatures, poor soil aeration, or vigorous competitive vegetation) or where a mixture of aspen and planted spruce are desired.     

Spatial point-pattern analysis for detecting density-dependent competition in a boreal chronosequence of Alberta

In the boreal forest of Alberta, fire and wind often open gaps in the canopy where late-successional species can establish and over time cause a shift in the species distribution from deciduous (e.g., trembling aspen) dominated to mixedwood, to shade-tolerant conifer (e.g., white spruce) dominated stands. This study attempted to understand the change of density-dependent competition in a boreal chronosequence and the role of tree competition in affecting stand structure and mortality. Four 1-ha stem-mapped plots were established to represent a chronosequence comprised of aspen dominated, mixedwood, and spruce dominated stands in Alberta. Second order spatial point-pattern analysis using Ripley's K(t) function showed that intraspecific competition is a prevailing force causing conspecific tree mortality and thus shaping the stand structure. The results of bivariate K(t) function analysis did not reveal sufficient evidence of interspecific competition. This suggested that competitive interaction among heterospecific trees was not strong enough to cause significant tree mortality, but the analysis of marked correlation function revealed that interspecific competition could have a negative impact on tree growth. This study highlights the importance of density-dependent competition in understanding stand dynamics of boreal forests over succession.     

Urea treatment reduced Heterobasidion annosum s.l. root rot in Picea abies after 15 years

Stump protection using chemical or biological agents is the main control measure against root and butt rot caused by Heterobasidion annosum s.l. in northern and temperate conifer forests. Long-term effects of urea treatment of stumps are poorly known and here we describe a 15-year study of urea treatment on the rot incidence on Norway spruce (Picea abies). We also tested the effect of urea protection on tree growth and on the resistance of stands against strong winds. Four treatments were made in two replications in two first-rotation P. abies stands in southern Sweden; after first-thinning stumps were (i) treated with urea 35% (w/v), (ii) artificially infected with H. annosum conidia, (iii) half urea treated and half artificially conidia infected, (iv) untreated, therefore naturally infected. After 15 years, the trees were sampled at 20 cm above ground using an increment borer and observed for presence of rot and, following incubation, presence of H. annosum conidia. Tree growth was calculated by measuring the diameter before and after the treatment. Urea treated plots showed the lowest incidence of rot (3%) as well as of H. annosum incidence (0%). Conidia treatment showed the highest incidence of rot (68%), its incidence was higher than that observed in natural infection treatment (43%), but did not differ from that of the 50% conidia treatment (47%). On about 30% of the rotted trees we observed conspicuous H. annosum fruiting. We did not observe growth reduction associated with tree rot. H. annosum was the only fungus observed associated with rotted trees which suggest that it was responsible for most of the rot observed in the investigated plots. Urea treated plots showed the lowest incidence of windthrown trees, and 59% of the windthrown tree incidence among the plots was explained by the incidence of H. annosum. Urea can be regarded as a reliable long-term protection method against root and butt rot of Norway spruce.     

Logging residue removal after thinning in Nordic boreal forests: Long-term impact on tree growth

The aim of this study was to determine the effect of whole-tree harvesting (WTH) on the growth of Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies (L.) Karst.) as compared to conventional stem harvesting (CH) over 10 and 20 years. Compensatory (WTH + CoF) and normal nitrogen-based (CH + F or WTH + F) fertilisation were also studied. A series of 22 field experiments were established during 1977-1987, representing a range of site types and climatic conditions in Finland, Norway and Sweden. The treatments were performed at the time of establishment and were repeated after 10-13 years at 11 experimental sites. Seven experiments were followed for 25 years.Volume increment was on average significantly lower after WTH than after CH in both 10-year periods in the spruce stands. In the pine stands thinned only once, the WTH induced growth reduction was significant during the second 10-year period, indicating a long-term response.Volume increment of pine stands was 4 and 8% and that of spruce stands 5 and 13% lower on the WTH plots than on CH during the first and the second 10-year period, respectively. For the second 10-year period the relative volume increment of the whole-tree harvested plots tended to be negatively correlated with the amount of logging residue. Accordingly, the relative volume increment decreased more, the more logging residue was harvested, stressing the importance of developing methods for leaving the nutrient-rich needles on site.If nutrient (N, P, K) losses with the removed logging residues were compensated with fertiliser (WTH + CoF), the volume increment was equal to that in the CH plots. Nitrogen (150-180 kg ha⁻¹) or N + P fertilisation increased tree growth in all experiments except in one very productive spruce stand. Pine stands fertilised only once had a normal positive growth response during the first 10-year period, on average 13 m³ ha⁻¹, followed by a negative response of 5 m³ ha⁻¹ during the second 10-year period. The fertilisation effect of WTH + F and WTH + CoF on basal area increment was both smaller and shorter than with CH + F.     

Impacts of hemlock looper defoliation on growth and survival of balsam fir, black spruce and white birch in Newfoundland, Canada

Hemlock looper (Lambdina fiscellaria fiscellaria (Guen.)) is an economically damaging defoliator that undergoes periodic outbreaks in Newfoundland, Canada. It defoliates and causes extensive tree mortality to its primary host, balsam fir (Abies balsamea [L.] Mill.). We quantified tree survival using data from permanent sample plots (PSPs) and growth reduction or release using dendrochronology, and related these impacts to defoliation severity determined from annual aerial defoliation survey data. Such impact relationships are necessary as a fundamental input to a Decision Support System. Growth and survival of balsam fir, black spruce (Picea mariana (Mill.) B.S.P.) and white birch (Betula papyrifera Marsh.) were assessed from 1996 to 2008 in 48 Newfoundland Forest Service PSPs, selected based on four classes of defoliation severity. Two years of severe (71-100%) defoliation resulted in almost complete mortality for balsam fir, 10 years after defoliation, whereas survival was 70-80% for black spruce and white birch. Lower defoliation severity (1-2 years of moderate (31-70%) or 1 year of severe) resulted in approximately 60% survival for balsam fir and no reduction in survival for black spruce and white birch. Maximum growth reduction of balsam fir was 10-15% with 1 year of moderate-severe defoliation, 35-40% with 2 years of moderate defoliation, and about 50% with 2 years of severe defoliation. Growth recovered to pre-defoliation rates 5 years after defoliation ceased in all severity classes. Growth reduction and recovery of black spruce were more variable and lower than for balsam fir, and white birch exhibited only minor (<10%) growth reduction during the defoliation year or 1 year after defoliation. Control measures should focus on avoiding severe defoliation for two consecutive years.     

Factors affecting the probability of windthrow at stand level as a result of Gudrun winter storm in southern Sweden

A storm in January 2005 in southern Sweden (Gudrun) caused large damage to forests, archaeological sites, buildings, power supply, telecommunications, and infrastructure. In this work, we analyzed the factors affecting the probability of windthrow at stand level (i.e. tree- and stand characteristics) in southern Sweden. We selected a sub-sample (1721 plots) of existing plots from the Swedish National Forest Inventory (inventoried in 2003 and 2004). To assess whether these plots had suffered damage or not, aerial photographs were taken in summer 2005. In order to analyze the effects of measured characteristics on damage, field data for each sub-sample plot were merged with the corresponding result of the aerial photo interpretation.We found that the plots damaged were typically well stocked mature stands, dominated by Norway spruce (Picea abies (L.) Karst.), and often also recently thinned. Probability of damage increased with increase in stand height. In mixed stands, decrease of proportion of Norway spruce decreased the probability of damage. An admixture of deciduous trees, leafless in this winter storm, showed larger effect in this sense than that of Scots pine.     

Nutrient cycling and soil leaching in eighteen pure and mixed stands of beech (Fagus sylvatica) and spruce (Picea abies)

Studies on the combined effects of beech–spruce mixtures are very rare. Hence, forest nutrition (soil, foliage) and nutrient fluxes via throughfall and soil solution were measured in adjacent stands of pure spruce, mixed spruce–beech and pure beech on three nutrient rich sites (Flysch) and three nutrient poor sites (Molasse) over a 2-year period. At low deposition rates (highest throughfall fluxes: 17 kg N ha⁻¹ year⁻¹ and 5 kg S ha⁻¹ year⁻¹) there was hardly any linkage between nutrient inputs and outputs. Element outputs were rather driven by internal N (mineralization, nitrification) and S (net mineralization of organic S compounds, desorption of historically deposited S) sources. Nitrate and sulfate seepage losses of spruce–beech mixtures were higher than expected from the corresponding single-species stands due to an unfavorable combination of spruce-similar soil solution concentrations coupled with beech-similar water fluxes on Flysch, while most processes on Molasse showed linear responses. Our data show that nutrient leaching through the soil is not simply a “wash through” but is mediated by a complex set of reactions within the plant–soil system.     

Persistent effects of fire severity on early successional forests in interior Alaska

There has been a recent increase in the frequency and extent of wildfires in interior Alaska, and this trend is predicted to continue under a warming climate. Although less well documented, corresponding increases in fire severity are expected. Previous research from boreal forests in Alaska and western Canada indicate that severe fire promotes the recruitment of deciduous tree species and decreases the relative abundance of black spruce (Picea mariana) immediately after fire. Here we extend these observations by (1) examining changes in patterns of aspen and spruce density and biomass that occurred during the first two decades of post-fire succession, and (2) comparing patterns of tree composition in relation to variations in post-fire organic layer depth in four burned black spruce forests in interior Alaska after 10-20 years of succession. We found that initial effects of fire severity on recruitment and establishment of aspen and black spruce were maintained by subsequent effects of organic layer depth and initial plant biomass on plant growth during post-fire succession. The proportional contribution of aspen (Populus tremuloides) to total stand biomass remained above 90% during the first and second decades of succession in severely burned sites, while in lightly burned sites the proportional contribution of aspen was reduced due to a 40-fold increase in spruce biomass in these sites. Relationships between organic layer depth and stem density and biomass were consistently negative for aspen, and positive or neutral for black spruce in all four burns. Our results suggest that initial effects of post-fire organic layer depths on deciduous recruitment are likely to translate into a prolonged phase of deciduous dominance during post-fire succession in severely burned stands. This shift in vegetation distribution has important implications for climate-albedo feedbacks, future fire regime, wildlife habitat quality and natural resources for indigenous subsistence activities in interior Alaska.     

Effects of soil moisture and species composition on growth and productivity of trembling aspen and white spruce in planted mixtures: 5-year results

Aspen (Populus tremuloides Michx.) and white spruce (Picea glauca (Moench.) Voss) were planted 0.5 m apart in intimate mixtures in 5 × 4 m plots, with two moisture regimes—irrigation versus control—and five species compositions—pure aspen (Aw100), mixed aspen and spruce (Aw83Sw17, Aw50Sw50, Aw17Sw83), and pure spruce (Sw100), replicated six times. Fifth-year assessments indicated that irrigation increased individual tree growth (height, RCD, crown width), plot leaf area index (LAI), and wood biomass. Increased aspen composition reduced the availability of soil moisture and consequently the growth of individual trees. With increased aspen composition more growth was allocated to stem in aspen and to foliage in white spruce. Comparatively, aspen responded more to irrigation and thus their growth is more dependent on precipitation than that of spruce. Among the three growth variables assessed, height responded more to irrigation in both species. Equal mixtures and aspen-dominated mixtures in control plots had higher productivity in terms of total wood biomass in both absolute and relative terms. The implications of these findings are discussed in relation to managing aspen and white spruce mixedwood forests under increasing drought expected as a result of climate change.     

Estimating “autotrophic” belowground respiration in spruce and beech forests: decreases following girdling

Seasonal fluxes of CO₂ from soil and the contribution of autotrophic (root + mycorrhizal) to total soil respiration (SR) were estimated for a mixed stand of European beech (Fagus sylvatica) and Norway spruce (Picea abies) in Central Europe. Mature trees of each species were girdled in August 2002 to eliminate carbohydrate allocation to roots. SR was measured at distances of 0.5, 1.0, and 1.5/2.0 m from the bole of each tree at 1–2 weeks intervals throughout the fall of 2002 and monthly during the spring and summer of 2003. The contribution of roots and mycorrhizae to total SR was estimated by the decrease in SR compared to ungirdled control trees to account for seasonal patterns evident in controls. SR decreased with soil temperature in the fall 2002 and increased again in 2003 as soil warmed. During most of the study period, SR was strongly related to soil temperature. During the dry summer of 2003, however, SR appeared to be uncoupled from temperature and was strongly related to soil water content (SWC). Mean rates of SR in beech and spruce control plots as well as root densities did not show a clear pattern with distance from the bole. SR decreased to levels below controls in beech within a few days after girdling, whereas spruce did not show a significant decrease until October 2002, 6 weeks after girdling. In both beech and spruce, decreased SR in response to girdling was greatest closest to the bole, possibly reflecting increased mycorrhizal activity close to the bole. Autotrophic respiration was estimated in beech to be as much as 50% of the total SR in the stand. The contribution of autotrophic respiration was less certain for spruce, although close to the bole, the autotrophic fraction may contribute to total SR as much as in beech. The large fraction of autotrophic respiration in total SR requires better understanding of tree level stresses that affect carbon allocation below ground.     

The nutrient status of Norway spruce in pure and in mixed-species stands

Atmospheric deposition of N and S appears to have caused nutrient imbalance in Norway spruce stands in southern Sweden. This calls for a change of forest management to procedures that promote nutrient balance. Studies have shown lower soil acidity in Norway spruce/deciduous mixed stands than in spruce monocultures, but the tree nutrient status in such mixtures has not been much investigated so far.

The nutrient status of Norway spruce foliage and top mineral soil chemistry in monocultures and in stands mixed with beech, birch, or oak was investigated through paired comparisons on 30 sites in southern Sweden (27 sites) and eastern Denmark (three sites). In total, 45 mixed stands and 34 pure stands were included in the study.

Spruce needles from mixed stands had higher concentrations and ratios to N of K, P, and Zn than needles from pure spruce stands. Among the mixed stands, the K status appeared to be positively correlated with the percentage of deciduous tree basal area. Soil samples from mixed stands had a higher Mg concentration, base saturation, and BC/Al ratio than soil samples from pure stands. The spruce needle nutrient status was comparable in pure stands on fertile sites and in mixed stands on poor sites. We did not detect any differences in spruce tree growth between pure and mixed stands.

This paper discusses possible reasons for a positive effect on the tree nutrient status in mixed-species stands and the possibility of using mixed-species stands as a forest management procedure to avoid nutrient imbalance.