Stocks and decay dynamics of above- and belowground coarse woody debris in managed Sitka spruce forests in Ireland

Coarse woody debris (CWD) has become recognised as an important component of the carbon (C) pool in forest ecosystems. In Ireland, managed Sitka spruce (Picea sitchensis (Bong) Carr.) forests account for 52.3% of the total forest estate. To determine the stock and decay dynamics of above and belowground CWD, field surveys using fixed area sample plots, were conducted in six even-aged Sitka spruce stands, representing the young, intermediate and mature stages of a typical commercial rotation. The volume, mass, density loss and C:N ratio of all CWD types (logs, stumps, and coarse roots) were determined using a five-decay class (DC) system. The decay rates and half life of CWD was also determined. To estimate CWD coarse root mass; roots associated with stumps classified in different decay classes were excavated. The coarse roots were categorised into small (2-10 mm), medium (10-50 mm) and large (>50 mm) diameter classes.CWD C-mass ranged from 6.98 to 18.62 Mg ha⁻¹ and was highest in an intermediate forest (D35), while the aboveground volume varied from 6.31 to 42.27 m³ ha⁻¹. Coarse roots accounted for 21% to 85% of the total CWD C-pool in the surveyed stands. The total CWD C-mass was poorly correlated with the number of thinning events (R² = 0.29), when data from D35 was excluded. The density loss was significant in logs (45%), stumps (58%), and small- (38%), medium- (50%) and large roots (38%) as decay progress from DC 0 to 4. There was a 46%, 41%, 51%, 72% and 57% decline in C:N ratio of logs, stumps, small-, medium- and large roots, respectively, as decay progressed from DC 0 to 4. The density decay rates were 0.059, 0.048 and 0.036 kg m⁻³ year⁻¹ for logs, stumps and coarse roots, respectively. The size classification of roots did not significantly affect their decay rate. The half life (50% decomposition) of CWD was estimated has 12-, 14- and 19 years for logs, stumps and roots of Sitka spruce. Regression curves showed a strong correlation between the density and C:N ratio (R² = 0.69, 0.74 and 0.93 for logs, stumps and coarse roots, respectively). The long term storage of C and its slow rate of decomposition make CWD a vital structural and functional component of the CWD C-pool and a major controller of forest ecosystem C-retention.     

Regeneration of Fitzroya cupressoides after indigenous and non-indigenous timber harvesting in southern Chilean forests

Fitzroya cupressoides (Cupressaceae) is an endemic and long-lived conifer of southern Chile and Argentina (40–43° S). This species has been subject to continuous exploitation since the 16th century, causing extensive population decline. Historically, the main labour force for the exploitation of F. cupressoides (alerce) was the indigenous Mapuche-Huilliche population, first under the command of the Spanish settlers and later, of non-indigenous Chileans. In coastal forests, timber of alerce has been harvested by Huilliche communities as well as by Chilean and international forestry companies. Records of the regeneration of this pioneer tree after exploitation in the Andean mountains have generally shown limited regeneration depending on the intensity of harvest. Because indigenous exploitation does not use machinery for timber harvesting, and is supposedly less utilitarian than commercial harvest, I propose that areas in the Coastal Range harvested by Huilliche communities should present higher regeneration of alerce than areas harvested by forestry companies. To test this hypothesis, I sampled 10 stands harvested and abandoned by forestry companies and 10 stands harvested by Huilliche communities in the coastal range of the Osorno Province (41° S, 400–800 m). In each stand, I estimated the density of regeneration (sapling stage), number of stumps, number of live adults, and the number of standing and fallen dead alerce. Each stand was characterized by elevation, forest-type, incidence of fire, and vegetation cover. Results showed that forests harvested by Huilliches had higher numbers of live, remnant adult trees with a dbh ≥ 60 cm than forests harvested by timber companies. The number of stumps with a dbh ≥60 cm was significantly higher in stands harvested by timber companies than in Huilliche stands. Despite large differences in sapling densities among stands, regeneration density of alerce was unrelated to the type of harvest used by indigenous people or forestry companies.     

A lack of large-diameter logs and snags characterises dead wood patterns in Irish forests

Dead wood is an important component of forest ecosystems and volumes vary depending on forest age, management intensity and productivity. This is the first large-scale study to quantify dead wood in Irish forests and to compare them to forests in other locations. We measured the volume and size distribution of logs, the density and size distribution of snags and the volume of dead wood contained in stumps in Oak (Quercus spp.) and Ash (Fraxinus excelsior) forests and in Sitka spruce (Picea sitchensis) plantations throughout Ireland. We also assigned each log, snag and stump to one of three decay classes (intact, part-rotted and well-rotted). We found no significant difference in log volume between any of the forest types. The majority (>90%) of logs were less than 20 cm in diameter, and large logs (>40 cm diameter) were scarce. We found a relatively high density of snags in all forest types but, as in the case of logs, over 90% of snags were <20 cm DBH and large snags (>40 cm DBH) were rare. The volume of dead wood contained in stumps was significantly higher in plantations than in Oak or Ash forests as a result of thinning and harvesting. Most logs and snags were moderately decayed but, in plantations, most stumps were intact. Log volume and the size of logs and snags were considerably lower than in old-growth forests in other regions. These patterns may reflect historical use of Irish forests for coppice and timber production. Management for biodiversity should aim to accelerate dead wood accumulation to increase the frequency of large-diameter logs and snags. Although management seeking to replicate the dead wood volumes of old-growth forests is ideal, it may be unrealistic in the short term.     

Bryophytes and decaying wood in Hepatica site-type boreo-nemoral Pinus sylvestris forests in Southern Estonia

The aim of the study was to establish the amount of decaying wood (logs and stumps) in various groups of Hepatica site-type pine forests of different age and management intensity and to analyse the composition of bryophytes in dependence of these factors. The average volume of CWD in old unmanaged forests was 47.5 m³/ha, which is rather well comparable with respective estimations from Fennoscandia. Reduced human impact contributes positively to the amount of CWD. Diversity of log diameter classes and decay stages is larger in old forests. Altogether 73 bryophyte species were recorded, 65 species on logs and 55 on stumps. Species richness on stumps was higher in managed forests than in unmanaged ones. At the same time, the species having high indicator value for man-cut stumps are very common species in boreal forests and grow on other substrata as well. Species composition and ecological conditions differed between stumps and logs. Logs are more humid microhabitats than stumps, therefore the occurrence of hepatics is more frequent on them. According to species composition on decaying wood the old unmanaged forests distinguished from others. As the differences of substratum characteristics were notable between old and young forests, the stand age described a considerable part of species variance on logs.     

Early seedling establishment of two tropical montane cloud forest tree species: The role of native and exotic grasses

Tropical montane cloud forest has been undergoing a drastic reduction because of its widespread conversion to pastures. Once these forests have been cleared exotic grasses are deliberately introduced for forage production. Exotic grass species commonly form monodominant stands and produce more biomass than native grass species, resulting in the inhibition of secondary succession and tree regeneration. The purpose of this study was to assess the effect of native vs. exotic grass species on the early establishment of two native tree seedlings (Mexican alder, Alnus acuminata and Jalapa oak, Quercus xalapensis) on an abandoned farm in central Veracruz, Mexico. Seedling survival and growth were monitored (over 46 weeks) in relation to grass cover and height, and available photosynthetic active radiation (PAR). More seedlings survived in the presence of the native grass Panicum glutinosum than those growing with the exotic grass Cynodon plectostachyus (92% vs. 48%). The causes of seedling mortality varied between species; Q. xalapensis was affected by herbivory by voles but mainly in the exotic grass-dominated stands, whereas A. acuminata seedlings died due to competition with the exotic grass. A. acuminata seedlings increased more in height in the exotic grass-dominated stands (102 ± 7.8 cm) compared to native grass-dominated stands (51 ± 4.7 cm). Grass layer height, cover and available PAR were correlated (Pearson; p < 0.05). In the exotic grass dominated plots, grass layer height was correlated with the relative height growth rates of Q. xalapensis (Pearson; p < 0.05). These results indicate that the exotic grass may be affecting tree regeneration directly (grass competition) and indirectly (higher herbivory). Passive restoration may occur once P. glutinosum dominated pastures are abandoned. However, when C. plectostachyus dominates, introduction of early and mid successional tree seedlings protected against vole damage is needed.     

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.     

Estimating the biomass and carbon pool of stump systems at a national scale

Countries that are signatories to the UNFCCC and its supplementary Kyoto Protocol are obliged to report changes in carbon pools. These should include the pool of carbon held in tree stumps and roots but, to date, few countries have been able to report this or separate it from the dead-wood pool. The aim of this study was to develop a general system for estimating and monitoring changes in stump system carbon using data from a traditional National Forest Inventory. The system was derived using data based on measurements of carbon (biomass) in inventoried permanent sample plots representing all relevant classes of land-use. With this design it was possible to trace matched carbon at the level of individual trees or stumps back to land-use prior to the 1990 baseline year. Between 1990 and 2003 in Sweden, the average annual net sink of stump systems was estimated to amount to 6.7 Mt CO₂ equiv. year⁻¹ – comparable to the reported net sink in 2008 of about 15 Mt CO₂ equiv. year⁻¹ from the whole Land Use, Land-Use Change and Forestry sector, which excluded any carbon in stump systems. In 2003 the carbon stock of stumps and roots was estimated at 495 Mt CO₂ equiv.; approximately five times that of the dead-wood pool as defined in Sweden, i.e. dead wood that mainly consists of boles. The Intergovernmental Panel on Climate Change requests that reported carbon should be matched to land-use and traced back to the 1990 base year; however, the present study confirms expectations that most carbon in stumps and roots is found on Forest land. The minimum requirements for estimating the carbon pool in stump systems at a national scale using the proposed methodology are that there should be: (i) a consistent time-series of harvest data, usually estimated as merchantable volume; (ii) conversion factors from merchantable volume to stump system biomass at death; and (iii) a representative decomposition model.     

Responses of polypore fungi following disturbance-emulating harvesting treatments and deadwood creation in boreal Norway spruce dominated forests

ABSTRACT

The emulation of natural disturbances in harvesting has become a widely accepted approach to reach ecologically sustainable forest management. The purpose of this study was to examine the responses of polypore fungi (species richness and composition) on four different cutting methods (selection, gap, patch, and clear-cutting) plus uncut controls, and the creation of high stumps and cut logs in combination with harvesting. Polypores were inventoried one year before, and two and seven years after cutting. A total of 71 polypore species were found, but no red-listed species were recorded seven years after cutting. Irrespective of the treatment, the variation in polypore richness was best explained by the number of deadwood objects in the stands. Species richness per deadwood object was the highest on natural logs. Species richness per volume of spruce deadwood was the highest on small-diameter deadwood and stumps, intermediate on natural logs and lower on cut logs and high stumps. These results indicate that cut logs created in harvesting cannot compensate the loss of deadwood in cutting and energy wood harvesting. Our results demonstrate the difficulty in rapidly creating habitats for red-listed polypores in commercially managed forests.     

Prescribed fire effects on bark beetle activity and tree mortality in southwestern ponderosa pine forests

Prescribed fire is an important tool in the management of ponderosa pine (Pinus ponderosa Dougl. ex Laws.) forests, yet effects on bark beetle (Coleoptera: Curculionidae, Scolytinae) activity and tree mortality are poorly understood in the southwestern U.S. We compared bark beetle attacks and tree mortality between paired prescribed-burned and unburned stands at each of four sites in Arizona and New Mexico for three growing seasons after burning (2004–2006). Prescribed burns increased bark beetle attacks on ponderosa pine over the first three post-fire years from 1.5 to 13% of all trees, increased successful, lethal attacks on ponderosa pine from 0.4 to 7.6%, increased mortality of ponderosa pine from all causes from 0.6 to 8.4%, and increased mortality of all tree species with diameter at breast height >13 cm from 0.6 to 9.6%. On a per year basis, prescribed burns increased ponderosa pine mortality from 0.2% per year in unburned stands to 2.8% per year in burned stands. Mortality of ponderosa pine 3 years after burning was best described by a logistic regression model with total crown damage (crown scorch + crown consumption) and bark beetle attack rating (no, partial, or mass attack by bark beetles) as independent variables. Attacks by Dendroctonus spp. did not differ significantly over bole heights, whereas attacks by Ips spp. were greater on the upper bole compared with the lower bole. Three previously published logistic regression models of tree mortality, developed from fires in 1995–1996 in northern Arizona, were moderately successful in predicting broad patterns of tree mortality in our data. The influence of bark beetle attack rating on tree mortality was stronger for our data than for data from the 1995–1996 fires. Our results highlight canopy damage from fire as a strong and consistent predictor of post-fire mortality of ponderosa pine, and bark beetle attacks and bole char rating as less consistent predictors because of temporal variability in their relationship to mortality. The small increase in tree mortality and bark beetle attacks caused by prescribed burning should be acceptable to many forest managers and the public given the resulting reduction in surface fuel and risk of severe wildfire.     

The burning characteristics of southeastern oaks: Discriminating fire facilitators from fire impeders

In southeastern pine–oak ecosystems, ecological restoration targets oaks for removal by chemical, mechanical, burning, or combinations of treatments. Managers often pursue oak removal indiscriminately despite the poorly understood historical structure, cover, and ecological function within these ecosystems. Restoration treatments often cite the impediment that oak litter represents to prescribed fire spread and effectiveness. We evaluated the burning characteristics of eight southeastern Quercus spp. by burning collected litter under controlled conditions in a combustion chamber. Replicated burns consisted of 15 g of litter on a 35 cm × 35 cm grid of xylene-soaked cotton strings. Burning characteristics measured included maximum flame height (cm), flaming duration (s), smoldering duration (s), residual ash, and mass loss rate (g s⁻¹). We compared all 8 oaks using ANCOVA, with litterbed depth as a covariate. The oaks differed for all burning characteristics measured (P < 0.001). Rank comparisons placed Quercus stellata and Quercus laevis as the species with greatest fire intensity, sustainability, and consumability, equivalent in many measures to longleaf pine and other fire resisters. Quercus virginiana and Quercus hemisphaerica burned with the least intensity, sustainability, and consumability, burning similarly to sand pine and other fire evaders. These results show that oaks common to southeastern United States ecosystems have litter properties, similar to pines, which vary in their ability to sustain fire. Understanding the pyric properties of oak species also suggests that managers prioritize removal of species that hinder prescribed fire effectiveness for restoration of southeastern USA pine–oak ecosystems.     

Short- and long-term effects of thinning and prescribed fire on carbon stocks in ponderosa pine stands in northern Arizona

Euro-American logging practices, intensive grazing, and fire suppression have increased the amount of carbon that is stored in ponderosa pine (Pinus ponderosa Dougl. Ex Laws) forests in the southwestern United States. Current stand conditions leave these forests prone to high-intensity wildfire, which releases a pulse of carbon emissions and shifts carbon storage from live trees to standing dead trees and woody debris. Thinning and prescribed burning are commonly used to reduce the risk of intense wildfire, but also reduce on-site carbon stocks and release carbon to the atmosphere. This study quantified the impact of thinning on the carbon budgets of five ponderosa pine stands in northern Arizona, including the fossil fuels consumed during logging operations. We used the pre- and post-treatment data on carbon stocks and the Fire and Fuels Extension to the Forest Vegetation Simulator (FEE-FVS) to simulate the long-term effects of intense wildfire, thinning, and repeated prescribed burning on stand carbon storage.The mean total pre-treatment carbon stock, including above-ground live and dead trees, below-ground live and dead trees, and surface fuels across five sites was 74.58 Mg C ha⁻¹ and the post-treatment mean was 50.65 Mg C ha⁻¹ in the first post-treatment year. The mean total carbon release from slash burning, fossil fuels, and logs removed was 21.92 Mg C ha⁻¹. FEE-FVS simulations showed that thinning increased the mean canopy base height, decreased the mean crown bulk density, and increased the mean crowning index, and thus reduced the risk of high-intensity wildfire at all sites. Untreated stands that incurred wildfire once within the next 100 years or once within the next 50 years had greater mean net carbon storage after 100 years compared to treated stands that experienced prescribed fire every 10 years or every 20 years. Treated stands released greater amounts of carbon overall due to repeated prescribed fires, slash burning, and 100% of harvested logs being counted as carbon emissions because they were used for short-lived products. However, after 100 years treated stands stored more carbon in live trees and less carbon in dead trees and surface fuels than untreated stands burned by intense wildfire. The long-term net carbon storage of treated stands was similar or greater than untreated wildfire-burned stands only when a distinction was made between carbon stored in live and dead trees, carbon in logs was stored in long-lived products, and energy in logging slash substituted for fossil fuels.     

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.     

Ectomycorrhizal fungal communities of rehabilitated bauxite mines and adjacent,natural jarrah forest in Western Australia

Species richness and species composition of ectomycorrhizal (EM) fungi were compared among rehabilitated mine sites and unmined jarrah forest in southwest Western Australia. Species richness, measured in 50 m × 50 m plots, was high. In the wetter, western region, mean species richness per plot in 16-year-old rehabilitated mine sites (63.7 ± 2.5, n = 3) was similar to that of unmined jarrah forest (63.6 ± 9.6, n = 9). In the drier, eastern region, species richness in 12-year-old rehabilitated mine sites (40.3 ± 2.1, n = 3) approached that of nearby forest (52.4 ± 9.3, n = 9). Species composition was analysed by detrended correspondence analysis. Rehabilitated sites of similar age clustered together in the analysis and species composition was closer to the native jarrah forest in the older rehabilitated plots. In unmined forest, species composition of fungal communities in the wetter, western region was different from communities in the drier, eastern region.     

The role of disturbance severity and canopy closure on standing crop of understory plant species in ponderosa pine stands in northern Arizona,USA

Concerns about the long-term sustainability of overstocked dry conifer forests in western North America have provided impetus for treatments designed to enhance their productivity and native biodiversity. Dense forests are increasingly prone to large stand-replacing fires; yet, thinning and burning treatments, especially combined with other disturbances such as drought and grazing, may enhance populations of colonizing species, including a number of non-native species. Our study quantifies plant standing crop of major herbaceous species across contrasting stand structural types representing a range in disturbance severity in northern Arizona. The least disturbed unmanaged ponderosa pine stands had no non-native species, while non-native grasses constituted 7–11% of the understory plant standing crop in thinned and burned stands. Severely disturbed wildfire stands had a higher proportion of colonizing native species as well as non-native species than other structural types, and areas protected from grazing produced greater standing crop of native forbs compared to grazed unmanaged stands. Standing crop of understory plants in low basal area thinned and burned plots was similar to levels on wildfire plots, but was comprised of fewer non-native graminoids and native colonizing plants. Our results also indicate that size of canopy openings had a stronger influence on standing crop in low basal area plots, whereas tree density more strongly constrained understory plant standing crop in dense stands. These results imply that treatments resulting in clumped tree distribution and basal areas <10 m² ha⁻¹ will be more successful in restoring native understory plant biomass in dense stands. Multiple types and severity of disturbances, such as thinning, burning, grazing, and drought over short periods of time can create greater abundance of colonizing species. Spreading thinning and burning treatments over time may reduce the potential for non-native species colonization compared to immediately burning thinned stands.     

Forest structure,habitat and carbon benefits from thinning floodplain forests: Managing early stand density makes a difference

Forest ecosystems are increasingly expected to produce multiple goods and services, such as timber, biodiversity, water flows, and sequestered carbon. While many of these are not mutually exclusive, they cannot all be simultaneously maximised so that management compromise is inevitable. We used a 42-year dataset from a naturally regenerating floodplain forest of the river red gum (Eucalyptus camaldulensis) to investigate the effects of pre-commercial thinning on long-term patterns in habitat quality, forest structure and rates of carbon storage (i.e. standing aboveground carbon). Estimates of habitat quality were based on the density of hollow-bearing trees because hollows are ecologically important to many species of vertebrates and invertebrates in these forests. Thinning improved habitat value by producing 20 (±8) hollow-bearing trees per ha after 42 years, while the unthinned treatment produced none. Unthinned (highest density) stands were dominated by many slender trees, mostly <25 cm in diameter, whereas thinned stands produced negatively skewed size distributions with higher median and maximum stem diameters. Moderately thinned stands (560 trees ha⁻¹) had the highest aboveground carbon storage rate (4.1 t C year⁻¹) and the highest aboveground carbon stocks (200.2 ± 9.6 t C ha⁻¹) after 42 years, while the unthinned treatment had the lowest carbon storage rate (1.6 t C year⁻¹) and an intermediate level of aboveground standing carbon (165.1 ± 31.1 t C ha⁻¹). Our results highlight the importance of early stand density as a determinant of long-term forest structure, habitat quality and carbon storage rates. We recommend that thinning be considered as one component of a broader strategy for enhancing the structure, habitat value and aboveground carbon storage of developing floodplain forests.     

Armillaria ostoyae in managed coniferous forests in Kastamonu in Turkey

Although several Armillaria species have been reported in Turkey, there is little information about their ecology in Turkish forests. In this study, we investigated five forest stands, approximately 5–74 ha in size, in Kastamonu province in the Black Sea Region of Turkey for the presence of Armillaria species in stumps and logs. The stands were mixed Abies nordmanniana ssp. bornmülleriana and Pinus sylvestris forests managed using a selective cuttings system; the proportion of fir in the total number of stems and stumps ranged from 36 to 98%. Based on sequence analysis of the internal transcribed spacer and intergenic spacer regions of the rDNA, all rhizomorphs sampled from the stumps and logs were of Armillaria ostoyae. The size of the genets was estimated with random amplified microsatellites analysis of the isolates and ranged from single stumps to approximately 450 m². One to seven genets were found in each stand. These results indicate that the genets had arisen from spores and vegetative spread was limited on most sites.     

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.     

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.     

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.     

A continuum of dead wood with various habitat elements maintains the diversity of wood-inhabiting fungi in an old-growth boreal forest

Evidence on habitat associations of threatened wood-inhabiting species in boreal forests may contribute to a better understanding of their ecology and conservation needs. We examined the diversity of wood-inhabiting fungal communities in an old-growth boreal forest with high substrate availability and continuity based on repeated surveys of fruit bodies. The number of species in morphological and functional groups in relation to coarse woody debris (CWD) attributes was estimated with generalized linear models. Additionally, we calculated species interaction networks of CWD attributes and fungal species. The composition of fungal communities was analysed using a non-metric multidimensional scaling with subsequent environmental fitting. Old conifer (especially spruce) logs and large aspen logs with branches represented the most important substrates for the red-listed species and the indicator species of old-growth boreal forests. Among “dynamic” CWD attributes such as time since tree death, decay class and stage of epixylic succession, the latter was the most important indicator of diversity of all species and their morphological and functional groups. The interaction network provided evidence of the importance of tree species diversity for fungal diversity. The composition of fungal communities was tree species specific and related to dynamic attributes, bark cover and diameter of logs. Our results suggest the importance of a continuum of dead wood from different tree species with a variety of niches such as branches, exposed wood, fragmented and complete cover of bark and patches of epixylic vegetation to maintain the assemblages of wood-inhabiting fungi in an old-growth boreal forest.