Growth of white spruce underplanted beneath spaced and unspaced aspen stands in northeastern B.C.—10 year results

Establishing white spruce (Picea glauca (Moench) Voss) by planting it under established aspen (Populus tremuloides Michx.), stands has substantial potential as a technique for regenerating boreal mixedwood stands. The presence of an aspen overstory serves to ameliorate frost and winter injury problems and suppresses understory vegetation that may compete with white spruce. In this study we examine the growth of white spruce during the first 10 years after being planted underneath a 39 year-old stand of trembling aspen following thinning and fertilization. Results indicate successful establishment and reasonable growth rates of white spruce planted under thinned and unthinned aspen stands, even with aspen basal area of 51 m² ha⁻¹. Thinning of overstory aspen to 1000 or 2000 stems ha⁻¹ did not increase light reaching seedlings, but did result in improvements in light above the shrub layer and in diameter and height growth of the underplanted seedlings. However, these increases in growth of underplanted spruce may not justify the expense of thinnings. Fertilization of these stands prior to planting had no effect on spruce growth. Growth of spruce underplanted at this site near Fort Nelson was similar to that at two other stands near Dawson Creek, B.C.     

Regeneration of Populus nine years after variable retention harvest in boreal mixedwood forests

Aspen and balsam poplar regeneration from root suckers were assessed in boreal mixedwood forests nine years after logging in a variable retention experiment (EMEND Project—Ecosystem Management Emulating Natural Disturbance) located north of Peace River, Alberta, Canada. Five levels of retention of mature trees (2%, 10%, 20%, 50% or 75% of the original basal area) were applied in stands dominated by aspen, white spruce or mixtures of the two species. Basal area of aspen (or that of aspen plus balsam poplar combined) prior to logging strongly influenced sucker density of aspen (or aspen + balsam poplar combined) and in some cases their growth. Nine years after harvest there was a decline in sucker density and volume ha⁻¹ with increasing retention levels of aspen (or both poplars combined); sucker density declined by 50% when only 20% of the original basal area was left in the stand. Retaining mature spruce trees in the stand had little influence on the number of suckers but did affect their total volume ha⁻¹. Thus, we suggest that by knowing stand aspen and balsam poplar density prior to logging and varying levels of retention of aspen and balsam poplar or conifers at harvest, the density of Populus regeneration can be predicted by managers, thereby allowing them to create a range of mixedwood conditions.     

Effects of overstory retention and site preparation on growth of planted white spruce seedlings in deciduous and coniferous dominated boreal plains mixedwoods

Survival and growth of planted white spruce was assessed under partial harvest treatments and different site preparation techniques in mixedwood forests of two compositions prior to logging: deciduous dominated (d-dom) – primarily comprised of mature trembling aspen (Populus tremuloides Michx.) and coniferous dominated (c-dom) – primarily comprised of mature white spruce (Picea glauca (Moench) Voss). Levels of overstory retention were 0% (clearcut), 50% and 75% of original basal area, and site preparation techniques were inverted mounding, high speed mixing, scalping and control (no treatment). The survival and growth of white spruce were assessed seven years after planting. The experiment was established as a part of the Ecosystem Management Emulating Natural Disturbance (EMEND) experiment located in northern Alberta, Canada. In the c-dom, the 50% and 75% retention of overstory resulted in reduced growth and survival of white spruce seedlings compared to clearcuts. In contrast, in the d-dom, the seedlings performed best in sites that had 50% of the overstory retained. For the c-dom, the mounding and mixing treatments yielded the best growth of spruce seedlings, while scalping yielded the worst. In the d-dom, spruce growth was highest in sites with the mixing treatment. In the d-dom, growth and survival of the planted spruce was greater than in the c-dom. The natural regeneration of deciduous trees was suppressed by the retention of canopy regardless of original composition.     

Carbon storage in old-growth and second growth fire-dependent western larch (Larix occidentalis Nutt.) forests of the Inland Northwest,USA

There is limited understanding of the carbon (C) storage capacity and overall ecological structure of old-growth forests of western Montana, leaving little ability to evaluate the role of old-growth forests in regional C cycles and ecosystem level C storage capacity. To investigate the difference in C storage between equivalent stands of contrasting age classes and management histories, we surveyed paired old-growth and second growth western larch (Larix occidentalis Nutt)–Douglas-fir (Pseudostuga menziesii var. glauca) stands in northwestern Montana. The specific objectives of this study were to: (1) estimate ecosystem C of old-growth and second growth western larch stands; (2) compare C storage of paired old-growth–second growth stands; and (3) assess differences in ecosystem function and structure between the two age classes, specifically measuring C associated with mineral soil, forest floor, coarse woody debris (CWD), understory, and overstory, as well as overall structure of vegetation. Stands were surveyed using a modified USFS FIA protocol, focusing on ecological components related to soil, forest floor, and overstory C. All downed wood, forest floor, and soil samples were then analyzed for total C and total nitrogen (N). Total ecosystem C in the old-growth forests was significantly greater than that in second growth forests, storing over 3 times the C. Average total mineral soil C was not significantly different in second growth stands compared to old-growth stands; however, total C of the forest floor was significantly greater in old-growth (23.8 Mg ha⁻¹) compared to second growth stands (4.9 Mg ha⁻¹). Overstory and coarse root biomass held the greatest differences in ecosystem C between the two stand types (old-growth, second growth), with nearly 7 times more C in old-growth trees than trees found on second growth stands (144.2 Mg ha⁻¹ vs. 23.8 Mg ha⁻¹). Total CWD on old-growth stands accounted for almost 19 times more C than CWD found in second growth stands. Soil bulk density was also significantly higher on second growth stands some 30+ years after harvest, demonstrating long-term impacts of harvest on soil. Results suggest ecological components specific to old-growth western larch forests, such as coarse root biomass, large amounts of CWD, and a thick forest floor layer are important contributors to long-term C storage within these ecosystems. This, combined with functional implications of contrasts in C distribution and dynamics, suggest that old-growth western larch/Douglas-fir forests are both functionally and structurally distinctive from their second growth counterparts.     

Aboveground biomass and nutrient accumulation dynamics in young black alder, silver birch and Scots pine plantations on reclaimed oil shale mining areas in Estonia

The growth, aboveground biomass production and nutrient accumulation in black alder (Alnus glutinosa (L.) Gaertn.), silver birch (Betula pendula Roth.) and Scots pine (Pinus sylvestris L.) plantations during 7 years after planting were investigated on reclaimed oil shale mining areas in Northeast Estonia with the aim to assess the suitability of the studied species for the reclamation of post-mining areas. The present study revealed changes in soil properties with increasing stand age. Soil pH and P concentration decreased and soil N concentration increased with stand age. The largest height and diameter of trees, aboveground biomass and current annual production occurred in the black alder stands. In the 7-year-old stands the aboveground biomass of black alder (2100 trees ha⁻¹) was 2563 kg ha⁻¹, in silver birch (1017 trees ha⁻¹) and Scots pine (3042 trees ha⁻¹) stands respective figures were 161 and 1899 kg ha⁻¹. The largest amounts of N, P, K accumulated in the aboveground part were in black alder stands. In the 7th year, the amount of N accumulated in the aboveground biomass of black alder stand was 36.1 kg ha⁻¹, the amounts of P and K were 3.0 and 8.8 kg ha⁻¹, respectively. The larger amounts of nutrients in black alder plantations are related to the larger biomass of stands. The studied species used N and P with different efficiency for the production of a unit of biomass. Black alder and silver birch needed more N and P for biomass production, and Scots pine used nutrients most efficiently. The present study showed that during 7 years after planting, the survival and productivity of black alder were high. Therefore black alder is a promising tree species for the reclamation of oil shale post-mining areas.     

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.     

Microclimatic and spruce growth gradients adjacent to young aspen stands

Effective utilization of a patch or cluster approach to growing aspen-white spruce mixtures in the boreal forest requires an understanding of how the faster growing aspen patches influence microclimate and spruce growth in adjacent spruce patches. In this paper we examine, how young (11–13 year old) aspen (Populus tremuloides Michx) patches influence microclimatic conditions in adjacent openings and how these are reflected in the growth of white spruce (Picea glauca [Moench] Voss) on three boreal mixedwood sites in west-central Alberta, Canada. Light levels increased continuously across the aspen stand boundary and reached maximum levels within the opening, while soil moisture was highest near the edge of the opening and then decreased with distance into the aspen stand or into the adjacent opening north of the aspen stand. Light levels were reduced over a greater distance when moving north from aspen stand edges compared to openings located south of the young aspen. These young aspen stands provided growing season frost protection within one tree length from the edge. The growth of white spruce was positively related to initial crown size and either light levels or distance from the edge. Stem volume growth was best predicted by initial tree size and the light levels at the midcrown of seedlings.     

The influences of conifer succession, physiographic conditions and herbivory on quaking aspen regeneration after fire

Fire suppression over the last century has increased conifer expansion and dominance in aspen-conifer forests, which appears to be a driving force behind aspen decline in some areas. The primary objective of this study was to examine how increasing conifer dominance affects aspen regeneration vigor following the return of fire. The influence of physiographic features and herbivory on aspen regeneration vigor were also examined. The study was conducted in the Sanford fire complex located in the Dixie National Forest in southern Utah, USA, where more than 31,000 hectares burned in the summer of 2002. Seven years after the burn (at 66 locations) we measured aspen regeneration density and height as response variables and former stand composition and density (the burned trees were still standing), soil characteristics, slope, aspect and presence or absence of herbivory as independent variables. Aspen regeneration (root suckering) densities ranged from <500 to 228,000 stems/hectare with an average of 37,000 stems/hectare. Post-fire aspen regeneration density was most strongly correlated with pre-fire stand successional status (as measured by stand composition and species abundance), with percent conifer abundance (R² = −0.55) and overstory aspen density (R² = + 0.50) being the most important. Average aspen suckering densities ranged from approximately 60,000 stems/hectare in what were relatively pure aspen stands (>90% aspen) to less than 5000 stems/hectare in stands where conifer abundance was greater than 90%. Soil C, N, and P showed positive correlations (R² = 0.07 to 0.17) with aspen regeneration vigor, while soil texture had a relatively weak influence on sucker regeneration. Aspen regeneration densities were 15% lower on north facing aspects compared to east, west and south facing aspects with slope steepness showing no correlation with regeneration vigor. Regeneration density was significantly lower (8%) at sites with evidence of herbivory versus sites where herbivory was absent. Overall, the aspen regeneration response in the Sanford fire complex was strong despite high wildlife densities, which may be related to disturbance size. Where the maintenance of aspen is desired in the landscape we recommend promoting fire when the percentage of overstory conifer stems is greater than 80% or overstory aspen density is less than 200 overstory stems/hectare.     

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.     

Effect of species and spacing of fast-growing nurse trees on growth of an indigenous tree, Hopea odorata Roxb., in northeast Thailand

We tested the effects of species and spacing of nurse trees on the growth of Hopea odorata, a dipterocarp tree indigenous to Southeast Asia, in a two-storied forest management system in northeast Thailand. Eucalyptus camaldulensis, Acacia auriculiformis, and Senna siamea were planted as nurse trees in 1987 at spacings of 4 m × 8 m, 2 m × 8 m, 4 m × 4 m, and 2 m × 4 m in the Sakaerat Silvicultural Research Station of the Royal Forest Department, Thailand. Seedlings of H. odorata were planted in the nurse tree stands at a uniform spacing of 4 m × 4 m and in control plots (no nurse trees) in 1990. Stem numbers of some nurse trees were thinned by half in 1994. The stem diameter and height of all trees were measured annually until 1995 and again in 2007. The mean annual increment (MAI) in volume was estimated as 8.2–10.1 m³ ha⁻¹ year⁻¹ for E. camaldulensis and 0.9–1.2 m³ ha⁻¹ year⁻¹ for S. siamea, smaller than reported elsewhere. This suggests that the site properties were not suitable for them. The MAI of A. auriculiformis was 7.9–9.8 m³ ha⁻¹ year⁻¹, within the reported range. Survival rates of H. odorata in the S. siamea stands and the control plots decreased rapidly during the first 2 years but then stayed constant from 1992. In contrast, survival rates of H. odorata in the E. camaldulensis and A. auriculiformis stands were initially high (>70%), but then decreased after 1995. Stem diameter, tree height, and stand basal area of H. odorata were large in both the S. siamea stands and the control plots from then. The growth of H. odorata was largest in the 2 m × 8 m S. siamea stands. In contrast, it was restricted in the E. camaldulensis and A. auriculiformis stands owing to strong shading by their canopies. Thinning by 50% tended to facilitate the growth of H. odorata temporarily in the E. camaldulensis and A. auriculiformis stands. The stand basal areas of nurse trees and of H. odorata showed a trade-off. These results suggest that the growth of H. odorata was maximized in the S. siamea stands. We assume, however, that the growth of H. odorata could be improved even in the E. camaldulensis and A. auriculiformis stands by frequent or heavy thinning.     

Specific gravity responses of slash and loblolly pine following mid-rotation fertilization

Wood quality attributes were examined in six stands of slash pine (Pinus elliottii Engelm. var. elliottii) and loblolly pine (P. taeda L.) in the lower Coastal Plain of Georgia and Florida. Several plots comprised each stand, and each plot was divided so that it received three fertilizer treatments: a control treatment with herbaceous weed control at planting and brush control at mid-rotation only (control); 45 kg ha⁻¹ N + 56 kg ha⁻¹ P + herbaceous weed control at planting and 224 kg ha⁻¹ N + 45 kg ha⁻¹ P + brush control at mid-rotation (fertilizer with N at planting); and 56 kg ha⁻¹ P + herbaceous weed control at planting and 224 kg ha⁻¹ N + 45 kg ha⁻¹ P + brush control at mid-rotation (fertilizer without N at planting). Ring width, ring earlywood specific gravity (SG), ring latewood SG, whole ring SG, and ring percent latewood were measured on each of seven trees. Of these measurements, this study focused mainly on the properties related to SG. Examination of the rings showed that latewood SG was significantly lower in trees treated with fertilizers with and without N at planting in the two to three years following fertilization, but that latewood SG gradually returned to a level similar to the control. Fertilizer without N at planting may also have had a brief negative effect on earlywood SG following mid-rotation fertilization, but it was not as clear or lasting as the effect on latewood SG. Additionally, although slash and loblolly pine appear to differ in the developmental patterns of these SG properties, there were no significant differences in how these patterns interacted with treatment. This study demonstrated that fertilization treatments have similar short-term effects on the SG of slash and loblolly pines, particularly in latewood, but the trees will return to a SG pattern consistent with unfertilized trees within two or three years.     

Mixed-species plantations of Acacia mangium and Eucalyptus grandis in Brazil: 1. Growth dynamics and aboveground net primary production

Nitrogen fertilizer inputs increased sharply over the last decade in Brazilian eucalypt plantations. Due to the economic and potential environmental cost of fertilizers, mixed plantations with N-fixing species might be an attractive option to improve the long-term soil N status. A randomized block design was set up in southern Brazil, including a replacement series and an additive series design, as well as a nitrogen fertilization treatment. The development of mono-specific stands of Eucalyptus grandis (0A:100E) and Acacia mangium (100A:0E) was compared with mixed plantations in proportions of 1:1 (50A:50E), and other stands with different densities of acacia for the same density of eucalypts. The objective was to assess the effect of inter-specific interactions on the early development of the two species. Aboveground biomass was measured 6, 12, 18 and 30 months after planting, sampling 6–10 trees of each species per treatment at each age, and allometric equations were established in 0A:100E, 100A:0E, 50A:50E and 50A:100E. The height and basal area of E. grandis seedlings were enhanced by 12% and 30%, respectively by N fertilization at age 1 year. Inter-specific competition led to a stratified canopy, with suppression in acacia growth earlier for basal area than for height. The mean number of stems per acacia tree at 36 months after planting was significantly higher in pure stands (3.7), than in 50A:50E (2.7) and in the additive series (between 1.6 and 1.8). H/D ratios were highly sensitive to inter-tree competition for the two species. The suppressed acacia understorey in mixed-species stands did not influence biomass production and partitioning within eucalypts. This pattern led to biomass accumulation combining the two species in 50A:100E that was about 10% higher than in 0A:100E, from age 12 months onwards. Aboveground net primary production (ANPP) amounted to 25 Mg ha⁻¹ and 37 Mg ha⁻¹ from age 18 to 30 months in 100A:0E and 0A:100E, respectively. Acacia ANPP in 50A:100E amounted to 2 Mg ha⁻¹ over the same period, as a result of substantial inter-specific competition. An increment in biomass production in these very fast-growing eucalypt plantations was achieved introducing acacia as an understorey and not in the 50A:50E design, as observed in other studies.     

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.     

The Brazil Eucalyptus Potential Productivity Project: Influence of water,nutrients and stand uniformity on wood production

We examined the potential growth of clonal Eucalyptus plantations at eight locations across a 1000+ km gradient in Brazil by manipulating the supplies of nutrients and water, and altering the uniformity of tree sizes within plots. With no fertilization or irrigation, mean annual increments of stem wood were about 28% lower (16.2 Mg ha⁻¹ yr⁻¹, about 33 m³ ha⁻¹ yr⁻¹) than yields achieved with current operational rates of fertilization (22.6 Mg ha⁻¹ yr⁻¹, about 46 m³ ha⁻¹ yr⁻¹). Fertilization beyond current operational rates did not increase growth, whereas irrigation raised growth by about 30% (to 30.6 Mg ha⁻¹ yr⁻¹, about 62 m³ ha⁻¹ yr⁻¹). The potential biological productivity (current annual increment) of the plantations was about one-third greater than these values, if based only on the period after achieving full canopies. The biological potential productivity was even greater if based only on the full-canopy period during the wet season, indicating that the maximum biological productivity across the sites (with irrigation, during the wet season) would be about 42 Mg ha⁻¹ yr⁻¹ (83 m³ ha⁻¹ yr⁻¹). Stands with uniform structure (trees in plots planted in a single day) showed 13% greater growth than stands with higher heterogeneity of tree sizes (owing to a staggered planting time of up to 80 days). Higher water supply increased growth and also delayed by about 1 year the point where current annual increment and mean annual increment intersected, indicating opportunities for lengthening rotations for more productive treatments as well as the influence of year-to-year climate variations on optimal rotations periods. The growth response to treatments after canopy closure (mid-rotation) related well with full-rotation responses, offering an early opportunity for estimating whole-rotation yields. These results underscore the importance of resource supply, the efficiency of resource use, and stand uniformity in setting the bounds for productivity, and provide a baseline for evaluating the productivity achieved in operational plantations. The BEPP Project showed that water supply is the key resource determining levels of plantation productivity in Brazil. Future collaboration between scientists working on silviculture and genetics should lead to new insights on the mechanisms connecting water and growth, leading to improved matching of sites, clones, and silviculture.     

Natural development and regeneration of a Central European montane spruce forest

Montane Norway spruce forests of Central Europe have a very long tradition of use for timber production; however, recently there has been increasing concern for their role in maintaining biological diversity. This concern, coupled with recent severe windstorms that led to wide-spread bark beetle outbreaks, has brought the management of montane spruce forests to the forefront of public policy discussions in Central Europe. In order to shed light on the natural development and current structure of mature montane spruce forests, we established four 0.25 ha research plots in a semi-natural montane spruce forest in the Šumava Mountains (The Bohemian Forest), Czech Republic. We mapped all trees, extracted increment cores for age and growth-pattern analyses, and inventoried all current tree regeneration, including the substrates on which it was found. Stands were characterized by uni-modal tree diameter distributions and high basal areas (56.6 m² ha⁻¹ on average), indicating a natural transition from the stem exclusion phase towards the understory reinitiation phase. The stands showed largely single-cohort recruitment age structures, however, with recruitment spanning seven decades. Our analyses suggest that this cohort existed as advance regeneration prior to major disturbances in the late 1800s, which included post-bark beetle salvage logging. Spatial pattern analyses of living and dead stems combined, showed an increase in uniformity of living trees, pointing to the role of natural density-dependent mortality. However, past growth patterns and historical documentation suggest that low intensity canopy disturbances (wind and snow) also caused mortality and diversified canopy structure. Because the stands developed naturally over the past 120+ years and thus escaped thinning operations, high volumes of coarse woody debris (94 m³ ha⁻¹) and snag densities (546 stems ha⁻¹) have accrued. Advance spruce regeneration was quite abundant and existed primarily on deadwood substrates, even though these occupied only a small percent of stand area. Because of salvage logging in the late 1880s, these stands do not qualify, according to the traditional paradigm, as natural spruce forests. As a result, they are recently subject to active management practices including salvage logging that remove dead and dying trees. Given the importance of deadwood for forest regeneration and recovery from disturbance, as demonstrated in this study, we argue that dead wood removal may limit future natural regeneration in these stands. Thus, the purported benefits of removing dead and dying trees from semi-natural forests must be carefully weighed against the potential detrimental impacts on natural spruce forest regeneration and biodiversity.     

Understory vegetation in old-growth and second-growth Tsuga canadensis forests in western Massachusetts

We compared the understory communities (herbs, shrubs, and tree seedlings and saplings) of old-growth and second-growth eastern hemlock forests (Tsuga canadensis) in western Massachusetts, USA. Second-growth hemlock forests originated following clear-cut logging in the late 1800s and were 108–136 years old at the time of sampling. Old-growth hemlock forests contained total ground cover of herbaceous and shrub species that was approximately 4 times greater than in second-growth forests (4.02 ± 0.41%/m² versus 1.06 ± 0.47%/m²) and supported greater overall species richness and diversity. In addition, seedling and sapling densities were greater in old-growth stands compared to second-growth stands and the composition of these layers was positively correlated with overstory species composition (Mantel tests, r > 0.26, P < 0.05) highlighting the strong positive neighborhood effects in these systems. Ordination of study site understory species composition identified a strong gradient in community composition from second-growth to old-growth stands. Vector overlays of environmental and forest structural variables indicated that these gradients were related to differences in overstory tree density, nitrogen availability, and coarse woody debris characteristics among hemlock stands. These relationships suggest that differences in resource availability (e.g., light, moisture, and nutrients) and microhabitat heterogeneity between old-growth and second-growth stands were likely driving these compositional patterns. Interestingly, several common forest understory plants, including Aralia nudicaulis, Dryopteris intermedia, and Viburnum alnifolium, were significant indicator species for old-growth hemlock stands, highlighting the lasting legacy of past land use on the reestablishment and growth of these common species within second-growth areas. The return of old-growth understory conditions to these second-growth areas will largely be dependent on disturbance and self-thinning mediated changes in overstory structure, resource availability, and microhabitat heterogeneity.     

Compositional changes of forest-floor vegetation in young stands of Norway spruce as an effect of repeated fertilisation

Forestry practices that aim to increase biomass production may mitigate climate change through increased carbon sequestration and the potential of substituting fossil fuels with renewable biofuels. Fertilising young stands of Norway spruce in Sweden have shown to increase tree growth by more than 200%. Fertilisation, however, also has other effects on forest ecosystems. Here, we studied the response of the species composition of forest-floor vegetation to three different frequencies of fertilisation in young stands of Norway spruce. Fertiliser was applied every year, every second year or every third year. The total amount of N ranged from 425 kg ha⁻¹ to 625 kg ha⁻¹, in combination with P, K, Ca, Mg, S, Mn, Zi, B and Cu. The largest effects of the fertilisation were found among bryophytes and lichens, which lost substantial cover. Unexpectedly, Deschampsia flexuosa, commonly known to be favoured by fertilisation, was negatively affected. Species that increased in frequency were Oxalis acetosella, Brachythecium sp. and Plagiothecium sp. Decreased availability of light, as an indirect effect of fertilisation through increased tree canopy cover, was found to be the most important factor behind the change in species composition of vascular plants. The total cover of bryophytes, however, did not show any significant response to the changes in canopy cover, indicating that the effects seen in this group may be a result of more direct effects of the fertiliser. Few significant differences were found between the two most intensive fertilisation frequencies, although fertilisation every third year was often distinguished from both the control and the other fertilised treatments. Even though the effects at the stand level were substantial, the effects on biodiversity and function of ecosystems on a landscape or regional level need further investigation.     

Biomass functions and expansion factors in young Norway spruce (Picea abies [L.] Karst) trees

Roots, stems, branches and needles of 160 Norway spruce trees younger than 10 years were sampled in seven forest stands in central Slovakia in order to establish their biomass functions (BFs) and biomass expansion factors (BEFs). We tested three models for each biomass pool based on the stem base diameter, tree height and the two parameters combined. BEF values decreased for all spruce components with increasing height and diameter, which was most evident in very young trees under 1 m in height. In older trees, the values of BEFs did tend to stabilise at the height of 3–4 m. We subsequently used the BEFs to calculate dry biomass of the stands based on average stem base diameter and tree height. Total stand biomass grew with increasing age of the stands from about 1.0 Mg ha⁻¹ at 1.5 years to 44.3 Mg ha⁻¹ at 9.5 years. The proportion of stem and branch biomass was found to increase with age, while that of needles was fairly constant and the proportion of root biomass did decrease as the stands grew older.     

Is the productivity decline in Norway spruce following whole-tree harvesting in the final felling in boreal Sweden permanent or temporary?

The growing demand for renewable energy sources in Sweden has resulted in an increased use of forest biomass that now includes logging residues. However, concern has been raised that the moderate increase in biomass removal associated with whole-tree harvesting results in a significant increase in nutrient removal, which in turn has a negative effect on future forest growth. Productivity over 31 years in planted Norway spruce (Picea abies (L.) Karst) in northern Sweden following three different harvest intensities is reported from a field experiment with exceptionally large growth reductions following whole-tree harvest. The three harvest intensities were applied in a randomized block design with four blocks: (i) conventional stem-wood harvest up to a top diameter of 5 cm (CH); (ii) whole-tree harvest of all above-stump biomass (WTH); (iii) branch and stem harvest with needles left on site (BSH). Recovery rate of biomass was almost 100% and the logging residues left were evenly spread over the 25- by 25-m experimental plots. Stand growth was negatively affected by WTH: basal area after 31 years was significantly lower following WTH (10.5 m² ha⁻¹) as compared to CH (14.0 m² ha⁻¹, p = 0.005) and BSH (14.2 m² ha⁻¹p = 0.003). Annual height growth of a sub-sample of trees (10 undamaged trees per plot, or 40 per treatment) was used to estimate and compare long-term effects on site productivity. This showed that stand growth loss resulted from a significant but temporary reduction in site productivity on WTH plots over a 5-year period (years 8-12, 1984-1988). Nitrogen is the major growth-limiting nutrient in boreal Swedish forests, and the N-content of needles during that period suggests that the temporal reduction in site productivity (i.e., stand growth) was primarily due to increased nitrogen loss with WTH.     

Effects of red alder and paper birch competition on juvenile growth of three conifer species in southwestern British Columbia

We examined the effects of competition from red alder (Alnus rubra Bong.) and paper birch (Betula papyrifera Marsh.) on the growth of western redcedar (Thuja plicata Donn), western hemlock (Tsuga heterophylla Sarg.), and Douglas-fir (Pseudotsuga menziesii Franco) at a site near Maple Ridge, British Columbia, Canada. At this site, the three coniferous species and two broadleaf species had been planted in 1999 as part of a long-term experiment examining effects of broadleaf density on conifer growth. Red alder and paper birch were planted alone and as a 50:50 mixture at four densities (0, 277, 556, and 1150 stems ha⁻¹).