Benchmarks and predictors of coarse woody debris in native forests of eastern Australia

Fallen coarse woody debris (CWD) is critical to forest biodiversity and function. Few studies model factors that influence CWD availability, although such investigations are critically needed to inform sustainable forest management. We assess benchmark levels of CWD in unharvested native forests and those harvested for timber, across a range of forests in north‐east New South Wales, Australia. We found timber‐harvesting was the dominant driver of CWD, with almost double the count (pieces ha^?1) and volume (m³ ha^?1) of total CWD in selectively harvested than unharvested sites. This pattern was consistent across wet and dry forest types. Harvested sites had greater counts of hollow‐bearing logs, and greater volumes of small and medium‐sized CWD (15–50 cm diameter) than unharvested sites. There was no effect of harvesting on the volume of large CWD (>51 cm diameter). Total volumes of CWD (>15 cm diameter) varied from 114 to 166 m³ ha^?1. We found few differences in CWD counts and volumes between forest types, with grassy woodlands and forests containing less CWD than other dry and shrubby forest types, reflecting lower potential input rates. The CWD levels recorded here are similar to those recorded in dry and wet sclerophyll forests elsewhere in Australia and are typical of global estimates for ‘old growth’ forests. Using general linear models we captured up to 57% of the variation in CWD across sites, and found that timber harvesting, topography and the numbers of standing hollow‐bearing and dead trees were significant predictors of CWD. Values for unharvested forest provide a benchmark that could be used to inform retention guidelines for CWD in managed forests in this region. Further assessment of the effect of repeat timber harvesting is needed to fully understand its impact on CWD dynamics, especially if forest residues resulting from timber harvesting are removed from native forests for bioenergy production.     

Responses of eight boreal flat bug (Heteroptera: Aradidae) species to clear-cutting and forest fire

Boreal flat bugs include a high proportion of species that are considered negatively affected by forestry. Knowledge on the biology and habitat demands of individual species is generally limited. We examined the influence on flat bugs of stand-age and clear-cutting, comparing five classes of spruce stands. The five classes were: clear-cut, unthinned, and thinned (all three products of current clear-cutting forestry), mature managed and old-growth stands (these two had never been clear-cut). We also compared unburned and recently burned mature pine forest. Fire, but not stand age, had a pronounced effect on species richness and total abundance. Aradus depressus showed a significant association with older forest stands. Aradus betulae occurred only in clear-cuts and burned forest indicating that this species is favored by disturbance in general. Aradus lugubris, Aradus crenaticollis and Aradus brevicollis were found only in the burned forest. Aradus brevicollis has not previously been shown to be associated with fire.     

Aboveground Net Primary Production and Leaf-Area Index in Early Postfire Vegetation in Yellowstone National Park

Aboveground net primary production (ANPP) and leaf-area index (LAI) of lodgepole pine (Pinus contorta var. latifolia Engelm. ex Wats.) saplings and aboveground productivity of herbaceous vegetation components were determined 9 years after the 1988 fires in Yellowstone National Park (YNP). Measurements were made in four sites representing a wide range of early postfire vegetation present in YNP, including high-density lodgepole pine, low-density lodgepole pine, and two nonforest stands. LAI of the pine saplings and total ANPP (trees plus herbs) generally increased with increasing sapling density, from 0.002 m² m^{− 2} and 0.25 Mg ha^{− 1} year^{− 1} in the infertile nonforest stand (100 pine saplings ha^{− 1}) to 1.8 m² m^{− 2} and 4.01 Mg ha^{− 1} year^{− 1} in the high-density pine stand (62,800 saplings ha^{− 1}). Aboveground herbaceous productivity was not strongly correlated with sapling density, but appeared to be influenced by soil fertility. In the high-density pine stand, tree ANPP and LAI were within the lower range of values reported for similar mature coniferous forests. This finding suggests that at least some ecosystem processes (related to ANPP and LAI) may have nearly recovered after only 9 years of postfire succession, in at least some of the young forests developing after the 1988 Yellowstone fires. Received 7 April 1998; accepted 1 December 1998.     

From Models to Measurements: Comparing Downed Dead Wood Carbon Stock Estimates in the U.S. Forest Inventory

The inventory and monitoring of coarse woody debris (CWD) carbon (C) stocks is an essential component of any comprehensive National Greenhouse Gas Inventory (NGHGI). Due to the expense and difficulty associated with conducting field inventories of CWD pools, CWD C stocks are often modeled as a function of more commonly measured stand attributes such as live tree C density. In order to assess potential benefits of adopting a field-based inventory of CWD C stocks in lieu of the current model-based approach, a national inventory of downed dead wood C across the U.S. was compared to estimates calculated from models associated with the U.S.’s NGHGI and used in the USDA Forest Service, Forest Inventory and Analysis program. The model-based population estimate of C stocks for CWD (i.e., pieces and slash piles) in the conterminous U.S. was 9 percent (145.1 Tg) greater than the field-based estimate. The relatively small absolute difference was driven by contrasting results for each CWD component. The model-based population estimate of C stocks from CWD pieces was 17 percent (230.3 Tg) greater than the field-based estimate, while the model-based estimate of C stocks from CWD slash piles was 27 percent (85.2 Tg) smaller than the field-based estimate. In general, models overestimated the C density per-unit-area from slash piles early in stand development and underestimated the C density from CWD pieces in young stands. This resulted in significant differences in CWD C stocks by region and ownership. The disparity in estimates across spatial scales illustrates the complexity in estimating CWD C in a NGHGI. Based on the results of this study, it is suggested that the U.S. adopt field-based estimates of CWD C stocks as a component of its NGHGI to both reduce the uncertainty within the inventory and improve the sensitivity to potential management and climate change events.     

Effects of Forest Type and Disturbance on Diversity of Coarse Woody Debris in Boreal Forest

Coarse woody debris (CWD) volume and diversity are vital attributes of forest ecosystems. However, despite their importance, their long-term dynamics associated with fire- or logging-origin and overstory type have not been examined in boreal forest. We hypothesize that (1) CWD compositional diversity increases with stand development whereas CWD volume follows a U-shaped pattern. Furthermore, we attempted to test if (2) CWD volume and compositional diversity converge for postlogged and postfire stands through stand development, and (3) mixedwoods have more CWD volume and greater compositional diversity than conifer or broadleaf overstory types. We sampled 72 stands ranging in age from 7 to 201 years in fire-origin stands and 7–31 years in managed stands with conifer, mixedwood, and broadleaf overstory types in central boreal Canada. For fire-origin stands, snag volume was 100–260 m³/ha in 7-year-old stands, 5–20 m³/ha in 25-year-old stands, and 25–60 m³/ha in older stands; downed woody debris (DWD) volume decreased from 7 to 72–90 year-old stands, increased in 124- to 139-year-old stands, then either decreased or increased in 201-year-old stands depending on overstory type. CWD diversity increased from 25 to 124–139 year-old and plateaued, but in 7-year-old stands, CWD diversity was as high as that in the 124 and up year-old age classes. Logging resulted in a smaller amount and lower size variability of CWD in 7-year-old stands, with a larger portion being fast-decomposing Betula papyrifera. Most CWD characteristics had not converged by approximately 30 years since disturbance between the two stand origins. More diverse CWD occurred in mixedwoods, but conifer stands contained the greatest CWD volume except in 7 year-old postfire stands. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users. B. W. Brassard collected and analyzed data and wrote the paper. H. Y. H. Chen conceived and designed the study, analyzed data, and critiqued earlier drafts of the paper.     

Damaged forests provide an opportunity to mitigate climate change

British Columbia (BC) forests are estimated to have become a net carbon source in recent years due to tree death and decay caused primarily by mountain pine beetle (MPB) and related post‐harvest slash burning practices. BC forest biomass has also become a major source of wood pellets, exported primarily for bioenergy to Europe, although the sustainability and net carbon emissions of forest bioenergy in general are the subject of current debate. We simulated the temporal carbon balance of BC wood pellets against different reference scenarios for forests affected by MPB in the interior BC timber harvesting area using the Carbon Budget Model of the Canadian Forest Sector (CBM‐CFS3). We evaluated the carbon dynamics for different insect‐mortality levels, at the stand‐ and landscape level, taking into account carbon storage in the ecosystem, wood products and fossil fuel displacement. Our results indicate that current harvesting practices, in which slash is burnt and only sawdust used for pellet production, require between 20–25 years for beetle‐impacted pine and 37–39 years for spruce‐dominated systems to reach pre‐harvest carbon levels (i.e. break‐even) at the stand‐level. Using pellets made from logging slash to replace coal creates immediate net carbon benefits to the atmosphere of 17–21 tonnes C ha^?1, shortening these break‐even times by 9–20 years and resulting in an instant carbon break‐even level on stands most severely impacted by the beetle. Harvesting pine dominated sites for timber while using slash for bioenergy was also found to be more carbon beneficial than a protection reference scenario on both stand‐ and landscape level. However, harvesting stands exclusively for bioenergy resulted in a net carbon source unless the system contained a high proportion of dead trees (>85%). Systems with higher proportions of living trees provide a greater climate change mitigation if used for long lived wood products.     

Soil carbon dynamics in a Pinus massoniana plantation following clear-cutting and slash removal

Aims Slash removal is a common practice to prepare recently harvested sites for replanting. However, little is known about its impact on soil carbon (C) dynamics in subtropical plantations. This study evaluates the effects of burning versus manual slash removal site preparation treatments on soil organic carbon (SOC), soil respiration and soil microbial community structure in a Pinus massoniana plantation in southern China.Methods Three areas within a mature P. massoniana plantation were clearcut. Two months following harvesting, slash on one-half of each area was burned (BURN), whereas slash was manually removed (MANR) on the other portion. Slash removal treatments were also compared with adjacent uncut plantation areas (UNCUT). Soil samples, and soil respiration measurements were used to characterize soil properties and microbial communities following slash removal treatments. Important findings Mean soil respiration rates from the MANR and BURN treatments were 26% and 17% lower, respectively, than the UNCUT treatment over 1 year. The MANR and BURN treatment resulted in soils with 27% and 9% reduction in total phospholipid fatty acids (PLFAs) and 18% and 10% reduction in bacterial PLFAs, respectively, compared with the UNCUT treatment. However, no significant differences existed between slash removal treatments with respect to soil chemical properties, SOC chemical compositions, soil respiration and microbial communities; although PLFA patterns were notably different for the burned plots. Most factors affecting C dynamics and microbial communities were not sensitive to the differences imparted to the ecosystem due to manual slash removal or burning. Our results suggested that low-intensity burning after clear-cutting might have no significant effect on soil C pool and its dynamics compared with manual slash removal in subtropical plantations.     

Fire and site type effects on the long-term carbon and nitrogen balance in pristine Siberian Scots pine forests

Effects of fire and site type on carbon (C) and nitrogen (N) balances were determined by following the change of total and component C and N pools along four chronosequences of fire-prone Siberian Scots pine ecosystems. These differed in the mean return interval of surface fires (unburned – moderately burned, 40 years – heavily burned, 25 years) and site quality (lichen versus Vaccinium site type). Of the Vaccinium site type (higher site quality) only a moderately burned chronosequence was studied. A total of 22 even-aged stands were investigated with stand ages ranging from 2 to 383 years. The C balance was dominated by the opposing dynamics of coarse woody debris (CWD) and biomass and could be divided into three phases: (1) Young stands (up to 40 years)acted as a net source for C of 6-10 mol C m-2 year-1 because the previous generation CWD pool originating from stand-replacing crown fires decayed much faster than biomass increased. During this period the C pool in the unburned lichen type chronosequence decreased from 807 to 480 mol C m-2. (2) Middle aged stands (40-100 years) being in a stage of maximum biomass accumulation were a net sink of 8-10 mol C m-2 year-1. (3)Maturestands (100 to > 350 years) continued to sequester C at a lower rate (0.8-2.5mol C m-2 year-1). Differences in the rates of C sequestration during the two later phases could be explained by the complex interaction between surface fire regime and site type. Recurrent surface fires resulted in enhanced mortality and regularly redistributed C from the living to the CWD pool thereby lowering the rate of C sequestration. Site quality determined the potential to recover from disturbance by fire events. Differences in site type did not correlate with soil and total ecosystem N pool size. However, the N status of needles as well as the N pool of physiologically active tissue was highest in the stands of the Vaccinium type. The woody C pool (biomass + CWD) was sensitive to differences in surface fire regime and site type. It was lowest in the heavily burned lichen type chronosequence (297 ± 108 mol C m-2), intermediate in the unburned and moderately burned lichen type chronosequence (571 ± 179 mol C m-2) and highest in the moderately burned Vaccinium type chronosequence (810 ± 334 mol C m-2). In contrast, the total soil C pool (organic plus mineral layer down to a depth of 25 cm) was independent of stand age, surface fire regimeand site type and fluctuated around a value of 250 mol C m-2. The organic layer C pool oscillated in response to recurring surface fires and its C pool was dependent on time since fire increasing at a rate of about 1.5 mol C m-2 year-during the first 40 years and then reaching a plateau of 170 mol C m-2. The total ecosystem N pool was 7.4 ± 1.5 mol N m-2 on average of which only 25 % were stored in biomass or coarse woody debris. Total ecosystem N was independent of stand age, surface fire regime and site type. No correlation was found between total ecosystem C and N pools. Average total ecosystem C:N ratio was 114 ± 35 mol C mol N-1. A conceptual model illustrating how changes in the regime of stand-replacing crown fires and recurrent surface fires and changes in site quality interact in determining the long-term C balance in Siberian Scots pine forests is presented.     

Accumulation and Connectivity of Coarse Woody Debris in Partial Harvest and Unmanaged Relict Forests

When a tree dies, it continues to play an important ecological role within forests. Coarse woody debris (CWD), including standing deadwood (SDW) and downed deadwood (DDW), is an important functional component of forest ecosystems, particularly for many dispersal-limited saproxylic taxa and for metapopulation dynamics across landscapes. Processes, such as natural disturbance or management, modify forest composition and structure, thereby influencing CWD abundance and distribution. Many studies have compared older forests to forests managed with even-aged silvicultural systems and observed a prolonged period of low CWD occurrence after harvesting. With fine-scale spatial data, our study compares the long-term impacts of light partial harvesting on the CWD structure of eastern deciduous hardwood forests. We mapped and inventoried DDW and SDW using variable radius plots based on a 10 m×10 m grid throughout an unmanaged, structurally-complex relict forest and two nearby forests that were partially harvested over 46 years ago. The relict stand had significantly larger individual pieces and higher accumulations of DDW and SDW than both of the partially harvested stands. Connectivity of CWD was much higher in the relict stand, which had fewer, larger patches. Larger pieces and higher proportion of decay-resistant species (e.g. Quercus spp.) in the relict forest resulted in slower decomposition, greater accumulation and increased connectivity of CWD. Partial harvests, such that occur with selection forestry, are generally considered less disruptive of ecosystem services, but this study highlights the long-term impacts of even light partial harvests on CWD stocks and distribution. When planning harvesting events, forest managers should also consider alternative methods to ensure the sustainability of deadwood resources and function.     

Post-Fire Spatial Patterns of Soil Nitrogen Mineralization and Microbial Abundance

Stand-replacing fires influence soil nitrogen availability and microbial community composition, which may in turn mediate post-fire successional dynamics and nutrient cycling. However, fires create patchiness at both local and landscape scales and do not result in consistent patterns of ecological dynamics. The objectives of this study were to (1) quantify the spatial structure of microbial communities in forest stands recently affected by stand-replacing fire and (2) determine whether microbial variables aid predictions of in situ net nitrogen mineralization rates in recently burned stands. The study was conducted in lodgepole pine (Pinus contorta var. latifolia) and Engelmann spruce/subalpine fir (Picea engelmannii/Abies lasiocarpa) forest stands that burned during summer 2000 in Greater Yellowstone (Wyoming, USA). Using a fully probabilistic spatial process model and Bayesian kriging, the spatial structure of microbial lipid abundance and fungi-to-bacteria ratios were found to be spatially structured within plots two years following fire (for most plots, autocorrelation range varied from 1.5 to 10.5 m). Congruence of spatial patterns among microbial variables, in situ net N mineralization, and cover variables was evident. Stepwise regression resulted in significant models of in situ net N mineralization and included variables describing fungal and bacterial abundance, although explained variance was low (R²<0.29). Unraveling complex spatial patterns of nutrient cycling and the biotic factors that regulate it remains challenging but is critical for explaining post-fire ecosystem function, especially in Greater Yellowstone, which is projected to experience increased fire frequencies by mid 21^st Century.     

Slash Pile Burning Effects on Soil Biotic and Chemical Properties and Plant Establishment: Recommendations for Amelioration

Ponderosa pine forest restoration consists of thinning trees and reintroducing prescribed fire to reduce unnaturally high tree densities and fuel loads to restore ecosystem structure and function. A current issue in ponderosa pine restoration is what to do with the large quantity of slash that is created from thinning dense forest stands. Slash piling burning is currently the preferred method of slash removal because it allows land managers to burn large quantities of slash in a more controlled environment in comparison with broadcast burning slash. However burning slash piles is known to have adverse effects such as soil sterilization and exotic species establishment. This study investigated the effects of slash pile burning on soil biotic and chemical variables and early herbaceous succession on burned slash pile areas. Slash piles were created following tree thinning in two adjacent approximately 20‐ha ponderosa pine (Pinus ponderosa) restoration treatments in the Coconino National Forest near Flagstaff, Arizona. We selected 30 burned slash pile areas and sampled across a gradient of the burned piles for arbuscular mycorrhizal (AM) propagule densities, the soil seed bank, and soil chemical properties. In addition, we established five 1‐m² plots in each burned pile to quantify the effect of living soil (AM inoculum) and seeding amendments on early herbaceous succession in burned slash pile areas. The five treatments consisted of a control (no treatment), living soil (AM inoculum) amendment, sterilized soil (no AM inoculum) amendment, seed amendment, and a seed/soil (AM inoculum) amendment. Slash pile burning nearly eliminated populations of viable seeds and AM propagules and altered soil chemical properties. Amending scars with native seeds increased the cover of native forbs and grasses. Furthermore adding both seed and living soil more than doubled total native plant cover and decreased ruderal and exotic plant cover. These results indicate that seed/soil amendments that increase native forbs and grasses may enhance the rate of succession in burned slash pile areas by allowing these species to outcompete exotic and ruderal species also establishing at the site through natural regeneration.     

Genetic variation in postfire aspen seedlings in yellowstone national park

A rare episode of regeneration of aspen (Populus tremuloides Michx.) by seeds occurred in Yellowstone National Park (YNP), Wyoming, USA, following extensive fires that occurred in 1988. In 1997, we sampled 410 aspen seedlings from 23 local populations distributed widely across YNP to determine how genetic diversity varies with elevation, substrate, plant competition, ungulate browsing, and geographical location. We employed 132 randomly amplified polymorphic DNA (RAPD) markers based on six primers to show genetic relationships within and among the postfire aspen seedling populations. Measures of genetic variation, including estimates of percentage polymorphic loci, expected heterozygosity, and Nei's FST, indicated that most of the variation occurred within rather than among local populations. There was no indication of geographical differentiation among sampled populations based on hierarchal estimates of Nei's FST, neighbour-joining, or correlations between genetic distance and geographical distance. Even genetically distant populations shared nearly 90% of the same markers. Within plots, the amount of genetic variation decreased slightly in response to increased percentage vegetative cover, mean seedling basal diameter, and mean seedling height. Geological substrate, density of lodgepole pine (Pinus contorta var. latifolia Dougl.) seedlings, browsing intensity, and elevation were not significantly related to levels of genetic variation within the seedling plots. These data suggest that genetic variation and geographical structure among seedling populations may occur over time as the transition from seedling-dominated stands to clone-dominated stands occurs.     

Decomposition Patterns of Foliar Litter and Deadwood in Managed and Unmanaged Stands: A 13-Year Experiment in Boreal Mixedwoods

Litter decomposition is a major driver of carbon (C) and nitrogen (N) cycles in forest ecosystems and has major implications for C sequestration and nutrient availability. However, empirical information regarding long-term decomposition rates of foliage and wood remains rare. In this study, we assessed long-term C and N dynamics (12–13 years) during decomposition of foliage and wood for three boreal tree species, under a range of harvesting intensities and slash treatments. We used model selection based on the second-order Akaike’s Information Criterion to determine which decomposition model had the most support. The double-exponential model provided a good fit to C mass loss for foliage of trembling aspen, white spruce, and balsam fir, as well as aspen wood. These litters underwent a rapid initial phase of leaching and mineralisation, followed by a slow decomposition. In contrast, for spruce and fir wood, the single-exponential model had the most support. The long-term average decay rate of wood was faster than that of foliage for aspen, but not of conifers. However, we found no evidence that fir and spruce wood decomposed at slower rates than the recalcitrant fraction of their foliage. The critical C:N ratios, at which net N mineralisation began, were higher for wood than for foliage. Long-term decay rates following clear-cutting were either similar or faster than those observed in control stands, depending on litter material, tree species, and slash treatment. The critical C:N ratios were reached later and decreased for all conifer litters following stem-only clear-cutting, indicating increased N retention in harvested sites with high slash loads. Partial harvesting had weak effects on C and N dynamics of decaying litters. A comprehensive understanding of the long-term patterns and controls of C and N dynamics following forest disturbance would improve our ability to forecast the implications of forest harvesting for C sequestration and nutrient availability.     

Variable carbon losses from recurrent fires in drained tropical peatlands

Tropical peatland fires play a significant role in the context of global warming through emissions of substantial amounts of greenhouse gases. However, the state of knowledge on carbon loss from these fires is still poorly developed with few studies reporting the associated mass of peat consumed. Furthermore, spatial and temporal variations in burn depth have not been previously quantified. This study presents the first spatially explicit investigation of fire‐driven tropical peat loss and its variability. An extensive airborne Light Detection and Ranging data set was used to develop a prefire peat surface modelling methodology, enabling the spatially differentiated quantification of burned area depth over the entire burned area. We observe a strong interdependence between burned area depth, fire frequency and distance to drainage canals. For the first time, we show that relative burned area depth decreases over the first four fire events and is constant thereafter. Based on our results, we revise existing peat and carbon loss estimates for recurrent fires in drained tropical peatlands. We suggest values for the dry mass of peat fuel consumed that are 206 t ha⁻¹ for initial fires, reducing to 115 t ha⁻¹ for second, 69 t ha⁻¹ for third and 23 t ha⁻¹ for successive fires, which are 58–7% of the current IPCC Tier 1 default value for all fires. In our study area, this results in carbon losses of 114, 64, 38 and 13 t C ha⁻¹ for first to fourth fires, respectively. Furthermore, we show that with increasing proximity to drainage canals both burned area depth and the probability of recurrent fires increase and present equations explaining burned area depth as a function of distance to drainage canal. This improved knowledge enables a more accurate approach to emissions accounting and will support IPCC Tier 2 reporting of fire emissions.     

Impacts of retention felling on coarse woody debris (CWD) in mature boreal spruce forests in Finland

We studied the immediate changes in pre-treatment coarse woody debris (CWD) after retention felling and mechanical site preparation (scarification) in mature Picea abies-dominated boreal forest. Retention felling and scarification were hypothetised to affect the amount of CWD. The disturbance caused to CWD was assumed to depend on species and decay class. Logs were inventoried before fellings, after fellings, and after scarification, estimating the damage percentage for each log. After felling, 7.8% of the total pre-treatment volume of CWD was destroyed in the felled area. After scarification, the decline from pre-treatment volume was 67.6% in the felled area. The amount of CWD decreased also inside the retention tree groups; in the 1st post-treatment season, 4.6% was destroyed of the pre-treatment volume and 20% in the 2nd season. Of the retained trees, 40% were uprooted by the end of the 2nd season. If the majority of the initial downed CWD is destroyed by scarification, as our results show, we can estimate that since scarification became a widely used regeneration method in the 1960s, at least from 4 to 6 million m³ of CWD has already been destroyed in Finland. The role of CWD as a key element for biodiversity in boreal forest is generally accepted, which has led to retention of trees in fellings instead of clear felling. We suggest that at least as important as leaving trees in order to maintain continuum in CWD and species diversity is to preserve existing CWD in fellings over the regeneration phase. This can be done using less destructive harvesting methods, reduced use of scarification and placing retention tree groups in patches with high amounts of CWD.     

Effects of fire on landscape heterogeneity in Yellowstone National Park,Wyoming

Abstract. A map of burn severity resulting from the 1988 fires that occurred in Yellowstone National Park (YNP) was derived from Landsat Thematic Mapper (TM) imagery and used to assess the isolation of burned areas, the heterogeneity that resulted from fires burning under moderate and severe burning conditions, and the relationship between heterogeneity and fire size. The majority of severely burned areas were within close proximity (50 to 200 m) to unburned or lightly burned areas, suggesting that few burned sites are very far from potential sources of propagules for plant reestablishment. Fires that occurred under moderate burning conditions early during the 1988 fire season resulted in a lower proportion of crown fire than fires that occurred under severe burning conditions later in the season. Increased dominance and contagion of burn severity classes and a decrease in the edge: area ratio for later fires indicated a slightly more aggregated burn pattern compared to early fires. The proportion of burned area in different burn severity classes varied as a function of daily fire size. When daily area burned was relatively low, the proportion of burned area in each burn severity class varied widely. When daily burned area exceeded 1250 ha, the burned area contained about 50 % crown fire, 30 % severe surface burn, and 20 % light surface burn. Understanding the effect of fire on landscape heterogeneity is important because the kinds, amounts, and spatial distribution of burned and unburned areas may influence the reestablishment of plant species on burned sites.     

Variation in soil seed bank species composition of a dry coniferous forest: spatial scale and sampling considerations

In order to assess the effects of disturbance on soil seed bank spatial structure, variation in seed bank species composition was analyzed at two spatial scales in mature (undisturbed) and recently disturbed stands of Interior Douglas-fir forest in south-central British Columbia. Coarse-scale analysis among stands 10–100 km apart using Multi-Response Permutation Procedures showed that individual mature stands were mainly characterized by unique seed bank species compositions. Disturbance had an homogenizing effect on seed bank species composition at the coarse scale––there was less variability among stands following disturbance by both low and high severity fires, and by non-salvage logging. In contrast, finer scale analysis among sampling units c. 10 m apart showed that heterogeneity in seed bank species composition was greater on severely burned and logged sites, commensurate with greater levels of soil disturbance, than on lightly burned and undisturbed sites. Despite the high intensity of seed bank sampling (sixty 25 cm² soil samples from each of 16 sites), species-area curves leveled off only when infrequent species were removed from the dataset. The number of seed bank samples required to account for the common species ranged from 84 on severely burned sites, to 196 on undisturbed sites; more than the 240 samples collected per disturbance class in this study would have been required to account for the remaining infrequent species. Overall, this study highlights the importance of sampling intensively within multiple stands to capture the variation in species composition inherent to these dry coniferous forest soil seed banks.     

The dynamics of old-growth Pseudotsuga forests in the western Cascade Range,Oregon, USA

Size and age structure, spatial analysis, and disturbance history were used to analyse the population structures and regeneration patterns of 8 conifer stands in the central western Cascade Range, Oregon, USA. Variation in forest structure reflected the effects of frequent (20–50 yr) low-intensity fires and treefalls, infrequent (100–200 yr) localised, intense fires, and extensive fires that resulted in stand replacement (every ca 400 yr?).The amount of canopy removed and the size of openings formed by fires and treefalls were important determinants of subsequent forest establishment. Single or several species stands of Pseudotsuga and/or Abies procera, or mixed species stands of Pseudotsuga, Abies procera, Tsuga heterophylla and Abies amabilis established in openings where intense fires had removed most of the canopy trees over several ha. Multi-tiered and multi-aged stands, often containing 400–500 yr-old Pseudotsuga and variously-sized more or less even-aged patches of younger shade tolerant Tsuga heterophylla and/or Abies amabilis, occurred where lower-intensity fires did not kill all overstorey trees or where treefalls occurred after the initial fire.Current regeneration processes are influenced by overstorey composition, the availability and size of canopy openings, and the availability of substrates suitable for regeneration. Tsuga heterophylla and Abies amabilis established under Pseudotsuga menziesii and Abies procera canopies and in small canopy openings (<400 m²) created by windfalls, but rarely under Tsuga. Down logs and stumps were favoured establishment sites for Tsuga.The disturbance regime of fires of low-to moderate-intensity, windfalls, and occasional fires that result in extensive stand replacement contrasts with the pattern of infrequent, catastrophic disturbances proposed for other areas of the Pacific Northwest. Although fires at stand establishment commonly determine much of the composition, structure, and subsequent stand development, canopy replacement by shade tolerant species occurs as the different life histories of the species are expressed in response to various disturbances differing in intensity and frequency. Such a non-equilibrium view of vegetation change is consistent with many other fire-dominated forests of the western United States.     

Microbial biomass C and N,and mineralizable-N,in litter and mineral soil under Pinus radiata on a coastal sand: Influence of stand age and harvest management

Microbial biomass C and N, and anaerobically mineralizable-N, were measured in the litter and mineral soil (0–10 cm and 10–20 cm depth) of Pinus radiata plantations in two trials on a nitrogen-deficient coastal sand. The trials comprised (a) stands of different age (1 to 33 years), with five of the seven stands studied being second rotation, and (b) a harvest-management trial, with stands established after different harvesting treatments of the first-rotation trees and understorey development controlled by manual weeding and chemical sprays. The harvest-management stands were sampled in the fifth year after the second-rotation establishment.In the stands of different age, the levels of microbial biomass C and N, and also mineralizable-N, in the litter and mineral soil showed no relationship with tree age and were similar to those in the oldest (33 years) stands of P. radiata. In the harvesting trial, five years after establishment of the second rotation, levels of microbial N and mineralizable-N in the litter and mineral soil were generally lowest where whole trees and the original forest floor had been removed; they were higher in associated plots in which the original forest floor had been removed but fertilizer N was regularly applied. No marked differences were then found between the other harvest treatments, viz. whole-tree harvest, stem-only harvest with slash remaining on site, and stem-only harvest plus extra added slash materials. In each trial, levels of microbial C and N and mineralizable-N were closely related to total C, and especially total N, in 0–10 cm depth mineral soil, but not generally in litter. Respiratory measurements strongly suggest that the microbial populations in mineral soil had a high metabolic activity.On an area basis in the harvest-management trial, total tree N and microbial N in the litter and mineral soil were lowest in stands where the original forest floor had been removed. In this particular treatment, microbial N in the litter plus mineral soil (0–20 cm depth) after five years of second-rotation growth comprised 7.3% of the total ecosystem N; values in the other treatments ranged between 5.6 and 6.0%.Our results emphasise the importance of slash and litter, and probably volunteer shrubs and herbaceous under-storey species, in conserving pools of potentially available N during the early stages of tree development.     

Oak woodland restoration: testing the effects on biodiversity of mycetophilids in southern Sweden

The effect of harvesting biofuel and woodland restoration on biodiversity is debated. To evaluate the effects of partial cutting on more organism groups, we used pairwise experimental and undisturbed control stands in a large landscape. On average 26% of the basal area including 50–95% of the understorey was harvested at each of 15 oak-rich forest sites. Our earlier results of partial cutting suggested a positive short-term effect for vascular plants and beetles and no or minor negative effects for fungi. Here we analyse the response of mycetophilids (Diptera: Sciaroidea excl. Sciaridae), a neglected but species-rich insect group that was strongly disfavoured by clear-cutting in previous studies. Increased deadwood (slash) caused increase in the number of individuals of mycetophilids associated with wood or wood fungi. The rarefaction species numbers of mycetophilids declined after the treatment, but the absolute number of species was not affected. Our results indicate that a 25% harvest can be compatible with conservation of vascular plants, fungi, saproxylic and herbivorous beetles and mycetophilids in temperate hardwood stands of the type studied. However, more studies are needed to determine what level(s) of stand thinning can be tolerated by different taxa in landscapes with small fragmented conservation stands.