Structure of a Central-European mountain spruce old-growth forest with respect to historical development

This study examines the structural characteristics of the tree layer, dead wood, canopy openings, and regeneration patterns of a spruce old-growth forest in the Bohemian Forest, Czech Republic. An old-growth stand with minor human influence and a stand that was presumably logged about 200 years ago were analyzed and compared, as some forest managers considered the presumable human impact as a reason for salvage logging. Even though the stands differed in tree density, height and DBH structure, it was not possible to conclude whether it was due to management history or the environmental differences. The volume of dead wood also differed between the stands. There was about 142 and 83 m³ ha⁻¹ of dead wood in the old-growth stand and presumably logged stand, respectively. The amount of dead wood found in the old-growth stand was comparable with values reported from spruce old-growth stands across Central Europe. In both stands, many canopy trees were arranged in linear patterns, which was a result of spruce regeneration on nurse logs. This suggests that the origin and development of the stands were characterized by natural processes and during the past 200 years typical old-growth structural characteristics have already evolved.     

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.     

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.     

Medicinal plants in old-growth, degraded and re-growth forests of NW Pakistan

Many old-growth forest stands in northwest Pakistan have been structurally transformed as a consequence of logging and livestock grazing, some of which are thereafter left to secondary succession. These forests represent an important resource for local inhabitants who gather and sell medicinal plants as part of their livelihood. With this in mind, the main objectives of our study were: (1) to assess differences in the structure of the tree layer and the abundance of medicinal plants among differently transformed forests, (2) to evaluate the recovery potential of medicinal plants under re-growth forests, and (3) to assess relationships between tree stand structural characteristics and the occurrence of medicinal plants.The first step of the study involved creating an approximate map covering an area of 90 km² for five forest-use types (old-growth forest, forest degraded by logging, derived woodland, agroforest and re-growth forest). Fifteen plots per forest-use type were randomly allocated at altitudes ranging from 2200 m to 2400 m asl, within which the abundance of 10 locally important medicinal herb species was assessed.The study stands differed greatly in tree basal area, which was highest in old-growth forest (48 m² ha⁻¹), lowest in agroforest areas (6 m² ha⁻¹) and intermediate in re-growth forest (20 m² ha⁻¹). All ten medicinal plant species were encountered in old-growth and in re-growth forests, but only five of these species also occurred on agroforest plots. The mean coverage of study medicinal plants was highest in old-growth forest (7%), low in forest degraded by logging, derived woodland and agroforest (0.3-2%), and intermediate in re-growth forest (4%). The Jaccard abundance based similarity index indicates a considerable similarity (0.6) between re-growth and old growth forest for both trees and medicinal plants. The overall abundance of medicinal plants increased with increasing tree basal area and canopy cover. The abundance of some particular species decreased; however, the most sought-after medicinal species Bergenia ciliata, Valeriana jatamansi and Viola cancescens increased with tree basal area within specific forest-use type and also across forest-use types. In conclusion, our data suggest that anthropogenic forest degradation leads to a reduction in the abundance of economically viable medicinal plants for the study region. It is further indicated that this can be reversed if degraded forests are allowed to regenerate.     

Alternative silvicultural practices with variable retention to improve understory plant diversity conservation in southern Patagonian forests

Understory plants could can act as indicators of temperate forest sustainability, health and conservation status due to their importance in ecosystem function. Harvesting impacts on understory plant diversity depends on their intensity. Variable retention has been proposed to mitigate the harmful effects of timber harvesting, but its effectiveness remains unknown in southern Patagonian Nothofagus pumilio forests. The objectives of this study were to: (i) define a baseline of understory plant diversity in old-growth forests along a site quality gradient and under canopy gaps; (ii) evaluate stands with three different variable retention treatments compared to old-growth forests; and (iii) assess temporal changes during 4 years after harvesting (YAH). A 61 ha N. pumilio forest was selected. Understory plant (Dicotyledonae, Monocotyledonae and Pteridophyta) richness, cover (including woody debris and bare forest floor) and aboveground dry biomass were characterized in summer for 5 years. Before harvesting, baseline samples were conducted along a site quality gradient and outside/inside canopy gaps. Analyzed treatments include a control of old-growth forest (OGF) and three different harvesting treatments with variable retention: (i) dispersed retention (DR) of 30 m² ha⁻¹ (20-30% retention); (ii) aggregated retention (AR) with one aggregate per hectare and clear-cuts (28% retention); and (iii) combined dispersed and aggregated retention (DAR) with one aggregate per hectare and dispersed retention of 10-15 m² ha⁻¹ (40-50% retention). Data analyses included parametric and permutational ANOVAs, multivariate classification and ordinations.Before harvesting, 31 plant species were found, where richness, cover and biomass were directly related to site quality. The presence of canopy gaps did not have a significant impact on the measured variables. After harvesting, 20 new species appeared from adjacent associated environments (two from N. antarctica forests and 18 from grasslands and peatlands). At the stand level, understory values were higher in AR > DR > DAR > OGF. Most (81-95%) plant richness at baseline conditions was conserved in all treatments, where inside the aggregates understory remained similar to OGF. Combination of aggregated and dispersed retention (DAR) better limited exotic species introduction and protected sensitive species, improving conservation in harvested stands. Changes in understory variables were observed after the first YAH in all treatments; greater changes were observed in the harvested areas than in aggregates. Changes stabilized at the fourth YAH. As a conclusion, the location of retention aggregates should be selected to preserve species understory diversity of more speciose and diverse habitats or particularly uncommon stands. Implementation of different kinds (patterns and levels) of retention for improvement of biodiversity conservation in harvested forests should be included in timber and forest management planning.     

Canopy gaps and regeneration in old-growth Oriental beech (Fagus orientalis Lipsky) stands, northern Iran

Virgin beech Fagus orientalis forests in northern Iran provide a unique opportunity to study the disturbance regimes of forest ecosystems without human influence. The aim of this research was to describe characteristics of natural canopy gaps and gap area fraction as an environmental influence on the success of beech seedling establishment in mature beech stands. All canopy gaps and related forest parameters were measured within three 25 ha areas within the Gorazbon compartment of the University of Tehran’s Kheyrud Experimental Forest. An average of 3 gaps/ha occurred in the forest and gap sizes ranged from 19 to 1250 m² in size. The most frequent (58%) canopy gaps were <200 m². In total, canopy gaps covered 9.3% of the forest area. Gaps <400 m² in size were irregular in shape, but larger gaps did not differ significantly in shape from a circle. Most gaps (41%) were formed by a single tree-fall event and beech made up 63% of gap makers and 93% of gap fillers. Frequency and diversity of tree seedlings were not significantly correlated with gap size. The minimum gap size that contained at least one beech gap-filling sapling (<1.3 m tall) was 23.7 m². The median gap size containing at least one beech gap-filling sapling was 206 m² and the maximum size was 1808 m². The management implications from our study suggest that the creation of small and medium sized gaps in mixed beech forest should mimic natural disturbance regimes and provide suitable conditions for successful beech regeneration.     

Availability of standing trees for large cavity-nesting birds in the eastern boreal forest of Québec,Canada

Large cavity-nesting birds depend on large-diameter trees for suitable nest sites. The increased spatial extent of commercial timber harvesting is modifying forest structure across the land base and may thus compromise the availability of large trees at the landscape scale. In this study, our objectives were to (1) characterize the availability of large living and dead trees in old-growth stands dominated by different tree species and surficial deposits that encompass the range of natural cover types of eastern Québec's boreal forest; (2) analyze the distribution of trees among decay-classes; and (3) compare the availability of large trees in unharvested, remnant, and harvested stands for the entire range of decay-classes. A total of 116 line transects were distributed across unharvested forests, remnant linear forests, and cutblocks in cutover areas. Unharvested forest stands (black spruce [Picea mariana], balsam fir [Abies balsamea]–black spruce, balsam fir–white spruce [Picea glauca] and balsam fir) reflected a gradient of balsam fir dominance. The remnant forests selected were isolated for 5–15 years. Analyses were performed at two diameter cut-off values. Trees with DBH ≥20 cm were considered for availability of total trees whereas trees with DBH ≥30 cm were considered for availability of large trees. Forest stands comprised high proportions of standing dead trees (33% of all stems, 8% were large dead stems). Availability of total and large standing trees increased with the dominance of balsam fir in stands. Forest stands located on thick surficial deposits showed higher densities of large dead trees for every stand type suggesting a higher productivity on those sites. Availability of stems according to decay-classes showed a dome-shaped distribution with higher densities of snags in intermediate decay stages. However, for large stems, black spruce stands showed a significantly lower availability that was consistent across all decay-classes. In linear remnant forests, pure balsam fir stands were absent. Remnant stands thus showed a much lower availability in large trees when compared with unharvested balsam fir stands. Clearcuts had the lowest densities of dead trees across sampled stands. Current even-aged management practices clearly affect availability and recruitment of large trees, therefore forest-dwelling wildlife relying on these structures for breeding is likely to be affected by large-scale harvesting in coniferous boreal forests.     

Structural and environmental characterization of old-growth temperate rainforests of northern Chiloé Island,Chile: Regional and global relevance

Old-growth forests are ecologically relevant reservoirs of biodiversity and provide valuable and unique ecosystem functions in the landscape. However, what constitutes an old-growth stand is confusing because the definition depends largely on the forest type under study. Despite the ecological importance of old-growth temperate rainforests in southern Chile in comparison to other global forests, no attempts have been made to characterize them as a way to assess their structural variability. Here, we characterized old-growth stands of Valdivian and North Patagonian rain forest types located in Chiloé Island (Chile, 42°30′S) using inventory data from 23 permanent plots (0.1 ha each) located in rural landscapes and protected areas of northern Chiloé Island. For each stand, its age (average age of the oldest trees present in each stand) and disturbance regimes (evidence of recent human impact, e.g. cuttings or fires, and tree growth rates) were used as defining old-growth criteria. We characterized the structure (tree species richness, size-density distributions, vertical stratification and presence of snags) and floristic composition of each stand. Environmental variables (i.e. temperature, distance to coastline and elevation) were related to stand structure using multivariate constrained correspondence analysis. Old-growth forests were commonly characterized by (a) tree basal areas >80 m²/ha; (b) density of shade-tolerant tree species in the emergent and dominant canopy layer >36%; (c) higher tree species richness (>7 tree species) than successional stands; (d) presence of large canopy emergents (>80 cm dbh, >25 m tall); (e) high vertical heterogeneity; and (f) minimum stand ages older than 200 years. Old-growth forests showed a distinctive structural variability and floristic diversity influenced both by stand age and disturbance history of the stands. Structural variability was also related to environmental differences among sites (e.g. air temperature, distance to coastline, soil types). Old-growth forest features described here can offer a baseline for managers interested in maintaining and restoring old-growth forest structure in southern temperate rain forests.     

Factors controlling the abundance of lianas along an altitudinal transect of tropical forests in Ecuador

In the light of putatively increasing liana abundances in present-day tropical forests and a persistent lack of understanding of liana abundance patterns and the responsible factors, we attempt to identify the key factors controlling liana abundance along an altitudinal transect in NE Ecuador. At four elevational levels (500, 1000, 1500, and 2000 m), each represented by 10 plots of 400 m², the abundance and diameters of all lianas (dbh ≥ 1 cm) and trees (dbh ≥ 10 cm) were recorded in old-growth forest stands in the Sumaco Biosphere Reserve (SBR). Results were analysed with available data on soil chemical properties and canopy structure.     

Recovery process of a mountain forest after shifting cultivation in Northwestern Vietnam

The recovery process of fallow stands in the mountainous region of Northwestern Vietnam was studied, based on a chronosequence of 1–26-year-old secondary forests after intensive shifting cultivation. The number of species present in a 26-year-old secondary forest attained 49% of the 72 species present in an old-growth forest. Total stem density decreased gradually from 172,500 ha⁻¹ in a 3-year-old forest to 24,600 ha⁻¹ in the 26-year-old stand, but stem density of larger trees (diameter at breast height (D) ≥ 5 cm) increased from 60 ha⁻¹ in a 7-year-old to 960 ha⁻¹ in the 26-year-old forests, which was similar to that of an old-growth forest. Annual biomass increment of the 26-year-old stand was 4.2 Mg ha⁻¹ year⁻¹. A saturation curve was fitted to biomass accumulation in secondary forests. After an estimated time of 60 years, a secondary forest can achieve 80% of the biomass of old-growth forests (240 Mg ha⁻¹). Species diversity expressed by Shannon Index shows that it takes 60 years for a secondary forest in fallow to achieve a plant species diversity similar to that of old-growth forests.     

Short-term community-level response of arthropods to group selection with seed-tree retention in a northern hardwood forest

We examined the short-term effects of group-selection harvesting with seed-tree retention on ground-dwelling and bark-dwelling arthropod communities in a northern hardwood forest in the Upper Peninsula of Michigan. Arthropods were sampled in 16 group-selection openings and 8 closed canopy reference plots. Two opening sizes were examined—radii of 0.5 (320 ± 27 m², n = 8) and 1.0 (1217 ± 62 m², n = 8) times the mean canopy height (22 m). Each opening and reference plot was centered on a single Betula alleghaniensis Britt. (yellow birch). Ground-dwelling arthropods were sampled using pitfall traps that were opened for two 1-week periods (rounds 1 and 2), and bark-dwelling arthropods were sampled with sticky traps attached to the centrally located B. alleghaniensis trees. Family-level diversity of ground-dwelling arthropods was lower in reference plots than in the openings, but the only significant difference occurred during round 2, between the matrix and large openings (P < 0.01). During both sampling periods, the ground-dwelling community exhibited a distinct clustering of family-level composition along environmental gradients such as opening size. Families such as Staphylinidae (rove beetles) and Trombidiidae (red velvet mites) were not favored by higher canopy openness while families such as Acrididae (grasshoppers) and Lycosidae (wolf spiders) were captured more in openings than in the forest matrix. Landing rates of wood-boring insects such as Buprestidae (metallic wood-boring beetles) and Xiphydriidae (wood wasps) were significantly higher on seed trees in group-selection openings than in reference plots (P < 0.05). Our results suggest that integrating group-selection openings within northern hardwood forests can lead to an increase in the family-level diversity of ground-dwelling arthropods, at least in the short term. Furthermore, seed trees left in such openings may be more attractive to bark- and wood-boring insects, which warrants further investigation into the susceptibility of these seed trees to damage by certain insect pests.     

Can the impact of deer browsing on tree regeneration be mitigated by shelterwood cutting and strip clearcutting?

Silvicultural treatments creating large canopy openings failed to restore regeneration of balsam fir (Abies balsamea (L.) Mill.) due to browsing pressure from white-tailed deer (Odocoileus virginianus Zimmermann). Consequently, we tested two alternative silvicultural treatments aimed at improving balsam fir establishment on Anticosti Island (Québec, Canada). In 1998 and 1999, we set up shelterwood seed cutting using three harvest intensities (0, 25 and 40% of basal area) and strip clearcutting with scarification using three different strip widths (15, 30 and 45 m), both with fenced and unfenced regeneration plots, in balsam fir stands. After 8 years, shelterwood seed cutting did not allow the establishment of new balsam fir seedlings, nor the development of unbrowsed balsam fir seedlings. In the strip clearcutting, deer browsing suppressed growth of palatable species in all strip widths. This favoured the development of unpalatable species, especially white spruce (Picea glauca (Moench) Voss). Our study demonstrates that the use of silvicultural treatments alone is unlikely to restore balsam fir regeneration on Anticosti Island, as long as the deer population remains higher than 20 deer/km².     

A comparison of lidar,radar, and field measurements of canopy height in pine and hardwood forests of southeastern North America

Forest canopy height is essential information for many forest management activities and is a critical parameter in models of ecosystem processes. Several methods are available to measure canopy height from single-tree to regional and global scales, but the methods vary widely in their sensitivities, leading to different height estimates even for identical stands. We compare four technologies for estimating canopy height in pine and hardwood forests of the Piedmont region of North Carolina, USA: (1) digital elevation data from the global Shuttle Radar Topography Mission (SRTM) C-band radar interferometry, (2) X- and P-band radar interferometry from the recently developed airborne Geographic Synthetic Aperture Radar (GeoSAR) sensor, (3) small footprint lidar measurements (in pine only), and (4) field measurements acquired by in situ forest mensuration. Differences between measurements were smaller in pine than in hardwood forests, with biases ranging from 5.13 to 12.17 m in pine (1.60–13.77 m for lidar) compared to 6.60–15.28 m in hardwoods and RMSE from 8.40 to 14.21 m in pine (4.73–14.92 m for lidar) compared to 9.54–16.84 in hardwood. GeoSAR measurements of canopy height were among the most comparable measurements overall and showed potential for successful calibration, with R² = 0.87 in pine canopies and R² = 0.38 in hardwood canopies from simple linear regression. An improved calibration based on differential canopy penetration is presented and applied to SRTM measurements, resulting in canopy height estimates in pine forests with RMSE and standard error <4.00 m. Each of the remotely sensed methods studied produces reasonable and consistent depictions of canopy height that can be compared with data of similar provenance, but due to differences in underlying sensitivities between the methods, comparisons between measurements from various sources require cross-calibration and will be most useful at broad scales.     

Methods for studying treefall gaps: A review

As silvicultural objectives have changed over the last several decades, managers are increasingly designing cutting regimes that mimic natural disturbance with the hopes that such systems will restore forests to a more natural condition while optimizing harvest yield. Treefall gaps, canopy openings caused by the death of one or more trees, are the dominant form of disturbance in many forest systems worldwide. These gaps play an important role in forest ecology by helping to maintain bio- and pedo-diversity, influencing nutrient cycling, and preserving the uneven-age nature of late-successional forests. In gap literature, there are inconsistencies with regard to gap terminology, methods for identifying and studying gaps, and modeling gap disturbances. From the papers reviewed, the size of treefall gaps ranges widely from 10 to >5000 m²; we suggest that the maximum gap size should be set at 1000 m². Larger openings tend to have microclimates and return intervals significantly different than smaller treefall gaps. Two main definitions of treefall gaps exist: canopy gap: a ‘hole’ in the forest through all levels down to an average height of 2 m above ground and extended gap: canopy gap plus the area that extends to the bases of surrounding canopy trees. Although researchers have assumed a variety of gap shapes to simplify measuring gap size, gaps are often irregularly shaped and so we recommend that gap areas and shapes be determined from detailed field measurements. Gap age may be determined from tree ring analysis of released trees in or near the gap edge, the spacing of whorls on released saplings, or from decomposition of gap-making trees. Windthrow is the main cause of canopy gaps in a variety of ecosystems; other causes include insects, diseases, acidic deposition, drought, and climate change. Treefall-gap models have been developed to predict the following processes during gap making or infilling: (i) gap abundance, (ii) forest structure, (iii) spatial and temporal variations in light levels, (iv) canopy dynamics, and (v) soil nutrient and water regimes. We recommend a protocol for gap studies and identify future research topics.     

Nitrogen leaching in response to increased nitrogen inputs in subtropical monsoon forests in southern China

Dissolved inorganic nitrogen (DIN) (as ammonium nitrate) was applied monthly onto the forest floor of one old-growth forest (>400 years old, at levels of 50, 100 and 150 kg N ha⁻¹ yr⁻¹) and two young forests (both about 70 years old, at levels of 50 and 100 kg N ha⁻¹ yr⁻¹) over 3 years (2004–2006), to investigate how nitrogen (N) input influenced N leaching output, and if there were differences in N retention between the old-growth and the young forests in the subtropical monsoon region of southern China. The ambient throughfall inputs were 23–27 kg N ha⁻¹ yr⁻¹ in the young forests and 29–35 kg N ha⁻¹ yr⁻¹ in the old-growth forest. In the control plots without experimental N addition, a net N retention was observed in the young forests (on average 6–11 kg N ha⁻¹ yr⁻¹), but a net N loss occurred in the old-growth forest (−13 kg N ha⁻¹ yr⁻¹). Experimental N addition immediately increased DIN leaching in all three forests, with 25–66% of added N leached over the 3-year experiment. At the lowest level of N addition (50 kg N ha⁻¹ yr⁻¹), the percentage N loss was higher in the old-growth forest (66% of added N) than in the two young forests (38% and 26%). However, at higher levels of N addition (100 and 150 kg N ha⁻¹ yr⁻¹), the old-growth forest exhibited similar N losses (25–43%) to those in the young forests (28–43%). These results indicate that N retention is largely determined by the forest successional stages and the levels of N addition. Compared to most temperate forests studied in Europe and North America, N leaching loss in these seasonal monsoon subtropical forests occurred mainly in the rainy growing season, with measured N loss in leaching substantially higher under both ambient deposition and experimental N additions.     

Comparison of soil CO2 flux between uncleared and cleared windthrow areas in Estonia and Latvia

Storms can turn a great proportion of forests’ assimilation capacity into dead organic matter because of windthrow and thus its role as a carbon sink will be diminished for some time. However, little is known about the magnitude or extent to which storms affect carbon efflux. We compared soil CO₂ fluxes in wind-thrown forest stands with different time periods since a storm event, and with different management practices (deadwood cleared or left on-site). This study examined changes in soil CO₂ efflux in two windthrow areas in north-eastern Estonia and one area in north-western Latvia, which experienced severe wind storms in the summers of 2001, 2002 and 1967, respectively. We measured soil CO₂ fluxes in stands formerly dominated by Norway spruce (Picea abies L. Karst.) with total and partial canopy destruction (all trees or roughly half of the trees in stand damaged by storm), in harvested areas (material removed after the wind storm) and in control areas (no damage by wind). Removal of wind-damaged material decreased instantaneous CO₂ flux from the soil surface. The highest instantaneous fluxes were measured in areas with total and partial canopy destruction (0.67 g CO₂ m⁻² h⁻¹ in both cases) compared with fluxes in the control areas (0.51 g CO₂ m⁻² h⁻¹), in the new storm-damaged areas where the material was removed (0.57 g CO₂ m⁻² h⁻¹) and in the old storm-damaged area where wood was left on site (0.55 g CO₂ m⁻² h⁻¹). The only factor affecting soil CO₂ flux was location of the measuring collar (plastic collar with diameter 100 mm, height 50 mm) - either on undamaged forest ground or on the uprooted tree pit, where the mineral soil was exposed after disturbance. New wind-thrown stands where residues are left on site would most likely turn to sources of CO₂ for several years until forest regeneration reaches to substantial assimilation rates. New wind-thrown stands where residues are left on site would most likely tend to have elevated CO₂ fluxes for several years until forest regeneration reaches to substantial assimilation rates. However, forest managers might be concerned about the amounts of CO₂ immediately released into the atmosphere if the harvested logs are burned.     

Transpiration and hydraulic traits of old and regrowth eucalypt forest in southwestern Australia

We tested the hypothesis that overstorey of eucalypt forest dominated by tall, large diameter trees uses less water than regrowth stands in the high rainfall zone (>1100 mm year⁻¹) of the northern jarrah (Eucalyptus marginata) forest in southwestern Australia. We measured leaf area, cover, sapwood area and sapwood density at three paired old and regrowth stands. We also measured sapflow velocity at one paired stand (Dwellingup) from June 2007 to October 2008. Old stands had more basal area but less foliage cover, less leaf area and slightly thinner sapwood. The ratio of sapwood area to basal area decreased markedly as tree size increased. Sapwood area of the regrowth forest stands (6.6 ± 0.30 m² ha⁻¹) was nearly double that of the old stands (3.4 ± 0.17 m² ha⁻¹), despite larger basal area at the old stands. Leaf area index of the regrowth stands (2.1 ± 0.26) was only one-third larger than that at the old stands (1.5 ± 0.15); hence, the ratio of leaf area to sapwood area was larger in old stands than in regrowth stands (0.45 ± 0.022 m² cm⁻² versus 0.32 ± 0.045 m² cm⁻²). Our results are consistent with theories that trees have evolved to optimize carbon gain rather than maintain stomatal conductance. Neither sapwood density (540–650 kg m⁻³) nor sap velocity differed greatly between regrowth and old stands. At the old forest site, daily transpiration rose from 0.5 mm day⁻¹ in winter to 0.9 mm day⁻¹ in spring–summer, compared to 0.9 mm day⁻¹ and 1.8 mm day⁻¹ at the regrowth site. Annual water use by the overstorey trees was estimated to be ∼230 mm year⁻¹ for the old stand and ∼500 mm year⁻¹ at the regrowth stand, or 20% and 44% of annual rainfall. The overwhelming role of stand sapwood area in determining stand water use, combined with the marked changes in the ratio of sapwood area to basal area with tree age and size, suggest that stand overstorey structure can be managed to alter overstorey water use and catchment water yield. Silviculture to promote old-forest-like attributes may be a viable means of delivering multiple water and conservation benefits.     

Long-term spatial dynamics of Acer saccharum during a population explosion in an old-growth remnant forest in Illinois

Species richness and evenness have greatly declined in oak–hickory forests in the central hardwood region in the U.S.A. in the past 100 years due to the rapid population growth of Acer saccharum. This study used a 50-year record of spatial dynamics to examine how demographic processes, particularly recruitment, may have contributed to this increase in an old-growth forest remnant, Brownfield Woods, Urbana, Illinois, U.S.A. The impact of canopy disturbance, including the outbreak of Dutch elm disease, on this increase was also evaluated. Historical maps of trees (≥7.6 cm DBH) from 1951, 1988, and 2001 in a 180 m × 280 m area were used to develop a series of univariate Ripley's L(d) functions to study changes in spatial patterns of three size classes of A. saccharum over time. Bivariate Ripley's L(d) functions were also utilized to evaluate spatial associations between recruitment and canopy disturbance. Our results indicated that A. saccharum was aggregated at most spatial scales up to 80 m during 1951–2001. Such aggregation arose mainly from small individuals. Furthermore, newly recruited individuals were aggregated at multiple spatial scales, and were significantly associated with canopy disturbance in general, as well as gaps created by Ulmus trees killed by Dutch elm disease. The aggregation of the 1951 initial group of small individuals changed via mortality to a random distribution over time. The results indicate that tree deaths caused by disturbances of different scales and types were the main cause of increased recruitment of A. saccharum in Brownfield Woods. The occurrence of Dutch elm disease further accelerated its population increase. This study demonstrated a direct spatial link between recruitment of A. saccharum and disturbance, and provided a long-term case study of a population explosion.     

Accumulation of dead wood in abandoned beech (Fagus sylvatica L.) forests in northwestern Germany

Currently, there is much debate about what strategy is most suitable for increasing old-growth attributes in forests that have been managed intensively for wood production in the past. Passive restoration, i.e. cessation of forestry interventions, should be considered when the old-growth attributes desired can be restored within a feasible period of time.Our study focuses on standing and lying coarse dead wood (≥20 cm diameter) in beech-dominated forests in northwestern Germany. We analyzed monitoring data of 545 sample plots (sized 500-1000 m²) from 12 strict forest reserves (SFRs). The SFRs had been without forestry intervention for up to 28 years.Both, number of dead objects and volume of dead wood (m³ ha⁻¹) increased significantly with ongoing time since abandonment from forestry interventions. The mean amount doubled from 9 to 18 m³ ha⁻¹ within 10 years. The proportion of standing dead wood was about 40% of the total dead wood pool ≥20 cm diameter.With mixed linear modeling we showed that dead wood increased by a mean net rate of about 1 m³ ha⁻¹ a⁻¹. Therefore, after three decades critical values for restoring the dead wood pool could be reached. We hypothesized that the rate of dead wood input is mainly determined by disturbance driven tree mortality such as oak decline, bark beetle infestations and storms.A comparison with primeval forests or reserves abandoned more than 100 years ago showed that the SFRs studied are at the beginning of a long process of dead wood accumulation.Based on our results, the abandonment of forest activities in harvestable pure and mixed beech stands is an effective strategy for restoring the dead wood pool.     

Alternative silvicultural practices with variable retention improve bird conservation in managed South Patagonian forests

Alternative silvicultural approaches to timber management, such as regeneration treatments with different degrees of stand retention, may mitigate negative effects of clear-cutting or shelterwood cuts in forested ecosystems, including changes in old-growth forest bird communities. The aims of this work were: (a) to compare bird species richness and densities among different silvicultural designs with variable retention (dispersed and/or aggregated) and unmanaged primary forests, and (b) to assess temporal changes at community and species levels before and after treatments. A baseline avian survey was conducted prior to harvesting to evaluate canopy gap presence and forest stand site quality influences. Subsequent to harvesting, data on bird species richness and density were collected by point-count sampling during the summer season for 5 consecutive years (4 treatments × 5 years × 6 sampling points × 5 counts). Bird species richness and density (15 species and 9.2 individuals ha⁻¹) did not change significantly with forest site quality of the stands and canopy gap presence in unmanaged forests. However, both variables were significantly modified in managed forests, increasing over time to 18 species and reaching to 39 individuals ha⁻¹. Inside the aggregated retention, bird communities were more similar to unmanaged primary forests than those observed within the dispersed retention or in clear-cuts. Opting for a regeneration method with dispersed and aggregated retention has great potential for managing birds in Nothofagus pumilio forests. This method retained enough vegetation structure in a stand to permit the establishment of early successional birds (at least in dispersed retention), and to maintain the bird species of old-growth forests which could persisted in the retention aggregates.