Gap dynamics in the Western Carpathian mixed beech old-growth forests affected by spruce bark beetle outbreak

Understanding how species-specific disturbances affect the dynamics of mixed forests is becoming increasingly important due to rapidly changing disturbance regimes. This study estimated the effect of Norway spruce (Picea abies (L.) Karst.) mortality on the disturbance processes in two mixed beech stands of the Western Carpathians that were affected by a bark beetle outbreak. We evaluated the size distribution, fraction of canopy and expanded gaps, the characteristics of gapmakers and the contribution of different species to gap size. The measured canopy gap fraction was <5%, and most canopy gaps were small (<100 m²). Spruce was the most abundant gapmaker, and its share among gapmakers was 3–6 times higher than its share in the canopy. We found that the increase in spruce mortality due to the outbreak resulted in a fine-scale mortality pattern. However, spruce gapmakers did not contribute much to gap area size, as shown by a weak correlation between the number of spruce gapmakers and the area of expanded gaps. Diameter distribution of living versus recently dead trees showed that beech mortality occurred disproportionately in large size classes. However, dead spruce trees were equally frequent in all diameter classes, which means beetles did not exclusively attack larger trees in these stands during the outbreak. We conclude that spruce mortality may have influenced successional processes by giving a competitive advantage to two other species that were not affected by the outbreak, provided that a high deer browsing intensity does not hinder the regeneration of new seedlings.

     

Tree recruitment in gaps of various size,clearcuts and undisturbed mixed forest of interior British Columbia,Canada

Tree seedling recruitment was monitored after various types of logging in mixed conifer and deciduous forests of northern British Columbia, Canada. Predicting tree seedling recruitment after disturbance is fundamental to understanding forest dynamics and succession and is vital for forest management purposes. Seedling recruitment success in multi-species northern latitude forests varied as a function of mature tree canopy cover, gap size and position in a gap. Recruitment was abundant within canopy gaps across a wide range of gap sizes (20–5000 m²), but recruit numbers dropped off rapidly under the closed forest canopy and in the open conditions of clearcuts. Inside canopy gaps, recruitment was similar by gap position in small gaps (<300 m²) but, in these northern latitude forests, exhibited a trend of increasing density from the sunny north to shady south end of larger gaps. This was true for all tree species regardless of their shade tolerance ranking. There was no evidence of gap partitioning by any of the tree species during the regeneration phase suggesting that adaptation to the subtleties of gap size during early recruitment are not well developed in these tree species. Favorable locations for emergence and early establishment of germinants were less favorable for growth and survival of established seedlings, i.e. the regeneration niches in these forests were discordant. Tree abundance and species diversity appears to be controlled more by differentiation among growth and survival niches than by the regeneration niches. From the perspective of forest management, abundant natural regeneration of all the dominant tree species of these mixed-species forests can be obtained after partial cutting.     

Regeneration of Norway spruce in canopy gaps in Sphagnum-Myrtillus old-growth forests

Gap-associated spruce (Picea abies (L.) Karst.) regeneration in Sphagnum-Myrtillus stands of south taiga forests (Central Forest Biosphere reserve, Tver region, Russia) was studied to evaluate the role of different disturbances in spruce dynamics. Sampled gaps (n=70) ranged from 40 m² to 1.7 ha in size, and from 1 to 70 years since disturbance moment. Formation of gaps lead to increase in the number of stems per ha in all gap size classes (small: 40–200 m², medium: 200–3000 m², and large: >3000 m² gaps). Spruce was the most important species in gap refilling, although its role was not the same in different gap classes. The highest values of relative abundance (compared to other species) were recorded in small gaps, and much lower values – in middle and large gaps. However, as refilling of gaps proceeded, spruce showed rather active regeneration in middle and large gaps and partly regained its abundance in middle-age disturbances. In general, all types of gaps studied supported spruce regeneration into the forest canopy. Almost perfect correlation between predicted outcome of spruce dynamics in gaps and its current role in the canopy of Sphagnum-Myrtillus stands suggests a good adaptation of this species to the current disturbance regime and a steady state of the these forests.     

Harvesting intensity affects forest structure and composition in an upland Amazonian forest

Forest structure and floristic composition were studied in a series of 0.5 ha natural forest plots at four sites near Porto Trombetas in Pará State, Brazil, 11–12 years after being subjected to differing levels of above-ground biomass harvest and removal. In addition to undisturbed control plots, experimental treatments included: removal of most trees ≥45 cm DBH (low intensity harvest); removal of trees <20 and ≥60 cm DBH (moderate intensity harvest); clear-cutting (100% above-ground biomass removal). Post-harvest basal area growth generally increased with harvest intensity, and total basal areas for trees ≥5 cm DBH were, at the time of our study, 60% (in the clear-cut) to about 80% of those in the control plots. Biomass harvests stimulated recruitment and growth of residual trees, particularly in the smaller diameter classes, but had little effect on species richness for small trees, seedlings, vine, herbs, and grasses. Species richness for trees ≥15 cm DBH was greater in the control and low-intensity (74–75 species) than in the moderate intensity (47 species) and clear-cut (26 species) treatment plots. While the tree flora within all harvest treatments was broadly similar to the undisturbed (control) plots and included similar numbers of species of the major plant families typical of the surrounding forests, the more intensive harvest treatments, especially the clear-cut, were dominated by a higher proportion of short-lived, early successional tree species.     

Regeneration dynamics of Sitka spruce in artificially created forest gaps

This study examined the variation in the development of naturally regenerated and planted seedlings of Sitka spruce (Picea sitchensis (Bong.) Carr.) within gaps cut in a 32-year-old stand of the same species. The circular gaps were 20 m in diameter and designed to allow sunlight into only half of the gap floor at midsummer given the latitude of 56°45′N. Eight plots (8 m × 3 m) were laid out along a north–south transect through each gap (four within the gap and two each under the closed canopy north and south of the gap). Each plot was sub-divided and seedlings were planted into one part and the other part was left to naturally regenerate. In subsequent seasons, plots were further subdivided into ‘weed free’ and ‘vegetation left untouched’. Results showed that while the two central plots within the gaps had the highest value of canopy openness, the highest accumulated temperature and lowest soil moisture were recorded in plots that received direct sunlight. However, level of germination was significantly higher in the shaded area of the gap than in the part that received direct sunshine suggesting that higher moisture levels in shaded areas are important to successful germination. Minimal germination was recorded in the plots beneath the canopy. Seedling survival was significantly influenced by the influx of competing vegetation, but only in the part of the gaps that received direct sunlight. The success of Sitka spruce regeneration within gaps appears to depend on sufficient moisture and light to support regeneration and early growth, but not too much light to encourage the development of competing vegetation. The permanently shaded areas of the gaps appeared to offer ground conditions with sufficient moisture and light to ensure successful germination and early growth of seedlings, but without excessive competition from other vegetation.     

Structure of Quercus gambelii communities in the Lincoln National Forest,New Mexico,USA

The structure of eight Quercus gambelii (Gambel's oak) communities in the Lincoln National Forest, New Mexico, USA were examined. Belt transects were used to estimate the density and basal area of the trees and the density of juvenile woody plants. In addition, diameter size-class distributions of Q. gambelii were examined to determine community development. The communities were estimated to be 109–137 years old and mid to late-successional. Total tree density was 3586–6480 plants/ha, with Q. gambelii having a relative density of 94–100%. Total basal area was 20.1–42.0 m²/ha, with Q. gambelii relative basal area 82–100%. The density and basal area of all other species present was low. Quercus gambelii juvenile density ranged from 1760 to 9160 plants/ha. Juveniles of all other species found were zero to 847 plants/ha. Based on Weibull analyses, all of the diameter size-class distributions of Q. gambelii were unimodal. There were few or no individuals in the smallest (1–5 cm) diameter size-classes, suggesting that recruitment of small Q. gambelii plants into the adult population may be below the replacement rate for these stands. Pooled size-class distributions for the other species were non-normal with most individuals in the smallest diameter size-classes. Low light levels below the canopy, a lack of canopy gaps, or browsing by Cervus elaphus (elk = red deer) may be the primary causes of poor recruitment because there were large numbers of Q. gambelii juveniles, but these individuals are not entering the small-tree size-class in any of the communities.     

Tree and forest encroachment into fescue grasslands on the Cypress Hills plateau,southeast Alberta,Canada

Tree encroachment into rough fescue (Festuca campestris) grassland has been identified as an ecological concern on the Cypress Hills plateau in southeastern Alberta, Canada. A combination of field sampling (109 transects), a dendrochronological assessment (1361 trees), and a time series analysis of remotely sensed images from five different time periods (1950–2002) were used to determine the extent and rate of tree encroachment and forest development. Tree cover increased by 768 ha (～51%) between 1950 and 2007, representing 10% of the study area, from 1502 ha of pre-1950 forest. Post-1950 tree invasion also created an ingress zone of 750 ha (～10% of study area) based on field transects. Forest cover increased at a consistent rate of 14.3 ha/year. Lodgepole pine (Pinus contorta var. latifolia) was the most common tree species associated with encroachment. Invasion based on the number of established trees occurred at an exponential rate of 3.1%/year after 1890, with density increased by filling spaces adjacent to and between trees within the grassland vegetation. The rate of tree establishment increased to 4.4%/year after 1980, suggesting a change occurred in environmental conditions. Annual atmospheric temperatures increased 0.55 °C from 1929 to 2005 (P < 0.001), with winter (December–February) and spring (March–May) temperatures accounting for most of the increase (P < 0.001), whereas summer (June–August) temperatures slightly decreased (P < 0.050, 0.34 °C) and precipitation increased (P < 0.005, 30 mm). Cattle and elk (Cervus elaphus) grazing was not considered a primary factor for explaining tree encroachment. Based on multidimensional scaling, lodgepole pine establishment was associated with warmer spring temperatures and greater fall (October–November) precipitation. A landscape devoid of wildfires, combined with greater moisture availability, and a longer frost-free season is likely conducive to the sustained establishment of coniferous trees and forest development within the Cypress Hills fescue grassland ecosystem. Long-term conservation of the fescue grasslands could be possible by reintroducing fire.     

Fuel mass and stand structure after post-fire logging of a severely burned ponderosa pine forest in northeastern Oregon

Stand structure and fuel mass were measured before and after a post-fire logging operation conducted 2 years after the 1996 Summit Wildfire (Malheur National Forest), in a ponderosa pine-dominated forest in northeastern Oregon. Variables were measured both pre- and post-logging in four replicate units for each of three treatments [un-logged control, commercial harvest (most dead merchantable trees removed), fuel reduction harvest (most dead merchantable trees removed plus most dead trees >10 cm diameter)]. Post-fire logging resulted in a significant decrease in mean basal area, down to 46% pre-treatment level in commercial units, and down to 25% in fuel reduction units. Logging significantly reduced tree density, especially for the smallest (<22 cm diameter) and intermediate (23–41 cm) diameter classes. Fuel reduction units also had significantly fewer snags (dead trees >30 cm diameter—4 ha⁻¹), compared to both commercial (23 ha⁻¹) units and to un-logged controls (64 ha⁻¹) in the year following timber harvest. Logging did not change ladder height or tree species composition (% ponderosa pine, Douglas-fir and grand fir). Total woody fuel mass increased significantly in fuel reduction units when compared to controls, with the greatest difference among treatments occurring in the slash fuel (<7.6 cm diameter) component (mean of 6.2 Mg/ha for fuel reduction stands versus 1.3 Mg/ha for un-logged stands). Logging activity caused no change in the mass of the forest floor (litter or duff). Model projections of the fuel bed using the fire and fuels extension of the forest vegetation simulator (FVS–FFE) indicate that the disparity in slash fuel mass between fuel reduction and un-logged units would be sustained until about 15 years post-logging, but a re-burn of moderate intensity occurring during this time would likely kill all young trees, even in un-logged units, because of the influence of other components of the fuel bed, such as grasses and shrubs. Model projections of 1000-h fuels (woody fuels >7.6 cm diameter) indicate that standing structure in all stands would collapse quickly, with the result that un-logged stands would contain two- or three-fold greater masses at 25 and 50 years post-logging, leading to much higher consumption rates of fuel in the event of a re-burn in the same place. Variation in dead tree fall and decay rates did not change the relationship among treatments in 1000-h fuel loads, but changed the time at which treatment differences were projected to disappear. Despite treatment differences in heavy fuel accumulations over time however, FVS–FFE predicts no differences among treatments in mortality of young trees due to either moderate or high intensity fire occurring in the same place at 25, 50, or 100 years post-fire logging. The lack of a re-burn effect is in part due to the reliance on flame length as the primary mechanism leading to tree death in the fire effect models used by FVS–FFE. If tree death turns out to be caused more by root burning or cambial heating, the observed variations in 1000-h fuel loadings among treatments could be significant in the event of a future re-burn.     

New management options in chestnut coppices: an evaluation on ecological bases

The effects of long rotation periods and heavy low thinning on chestnut (Castanea sativa Mill.) coppices have been evaluated from a bio-ecological point of view. Aboveground tree biomass and its partitioning, bio-ecological indexes such as litter production, leaf area index (LAI), radiation regime, and growth efficiency have been analyzed for 4 years in thinned and unthinned permanent plots established in a coppice stand aged 11 years under normal rotation (NR) and in a coppice stand aged 35 years under long rotation (LR). A decrease in LAI, litter production and growth rate with the age was observed. LR showed high current annual increments (>16.0 m³ ha⁻¹ per year and 8.0 Mg ha⁻¹ per year for volume and aboveground biomass, respectively). Only slight differences in growth efficiency were observed. The adoption of heavy thinning (one third of basal area removal) affected stocking and determined significant differences in the light regime below the canopy, amount of gaps in canopy cover and LAI values, particularly in the years immediately after thinning, whereas slight differences were observed in growth efficiency. Nevertheless, chestnut showed a good aptitude, more evident in the younger stand, to re-build a homogeneous canopy cover: only a few years after thinning, canopy cover characteristics of thinned plots were similar to those of control plots and differences were not significant. Growth and increments in thinned plots were practically the same as in control plots, a consequence of consistently higher performances of released trees in the thinned plots. The results concur to a positive evaluation of a cultivation system based on long rotation periods and heavy thinnings, not only for the obtainable revenue, but especially from a bio-ecological point of view and make it a valid alternative either to abandonment or traditional over-exploitation of chestnut coppice stands.     

Gap-Disturbance Regimes in Different Forest Types of Japan

Abstract

In this study, I defined a gap as a small opening formed in a forest canopy (area < 0.1 ha) and tried to synthesize gap-disturbance regimes of primary mature stands in different forest types of Japan, such as warm temperate evergreen broad-leaved (4 stands in 3 localities), cool temperate deciduous broad-leaved (10 in 5) and subalpine evergreen coniferous (3 in 1) forests. Mean percentage of the total gap area within the total forested area (percentage gap area) in each forest type was 17.0% in warm temperate (number of surveyed gaps was 161), 13.8% in cool temperate (278 gaps), and 8.0% in subalpine (100 gaps) forests. Mean gap density (ha^-1) and mean gap size (m²) were 19.5 and 77.1 in warm temperate, 16.4 and 92.0 in cool temperate, and 19.1 and 41.9 in subalpine forests, respectively. These figures indicate that gap density is not substantially different among the forest types, but the mean gap size of subalpine forests is smaller than the other two, resulting in lower percentage gap area of this forest type. The gap size distributions were similar among the forest types; smaller gaps were much more frequent than larger ones, and gaps > 400 rrr were rare in warm temperate and cool temperate forests. In subalpine forests gaps > 200 m² were rare. Gaps formed by multiple gapmakers comprised 19.9% of all gaps in warm temperate, 9.9% in cool temperate, and 44.9% in subalpine forests, which implies that gap formation by simultaneous tree fall or gap enlargement is more frequent in subalpine forests. Canopy trees died less often by uprooting in every forest type; dominant mode for the death of canopy trees was by leaving standing-dead or with broken trunks in every forest type. Since typhoons are obviously a chief agent of forest disturbance in Japan, frequency or magnitude of typhoon disturbance may influence these differences in the gap-disturbance regimes. In addition to the disturbance, tree architectures seem to affect some of these differences; narrower crown size of conifers compared with broad-leaved trees is considered one major cause for smaller gap size of the conifer forest.     

Effect of spacing and water availability on root:shoot ratio in Eucalyptus camaldulensis

Our limited understanding of the processes that control the allocation of biomass in trees is one of the factors that hinders our ability to develop mechanistic models of tree growth. Furthermore, accurate assessment of carbon sequestration by forests is hampered by lack of information regarding below-ground biomass. Below-ground to above-ground biomass ratios (BGB:AGB) are known to vary with a number of environmental factors, tending to increase in drier, harsher conditions. However, there are few, good datasets of BGB:AGB ratios of large trees, especially native Australian species. We aimed to investigate the response of BGB:AGB to water availability and tree spacing in 10-year-old Eucalyptus camaldulensis growing in a plantation in a low rainfall area.We carefully harvested 16 trees, ranging in diameter at breast height (DBH) from 7.6 to 25 cm, from a research trial near Deniliquin, NSW. Four replicates of each treatment from a factorial design with wide (3 m × 6 m) and narrow (3 m × 1.5 m) spaced trees and with natural rainfall (408 mm/year) (control) or irrigated plots (flooded six times per year) were selected. Above-ground tree parts were harvested separating stem, branch and foliage. Soil cores to 1 m depth were taken to sample small roots (<15 mm diameter) within each plot, then all roots >15 mm belonging to each tree were excavated using compressed air and an excavator. Roots were separated into six size classes within the range from <2 to >50 mm.Both water and spacing treatments influenced tree growth with trees being larger in irrigated, wide spaced plots. The BGB:AGB ratio was strongly influenced by irrigation (0.68 control, 0.34 irrigated) but not spacing. Allometric analysis of above- and below-ground biomass as a function of DBH showed that the relationship between DBH and above-ground biomass was conserved across treatments.By contrast, the relationship between DBH and below-ground biomass was influenced by water availability, commensurate with the large differences in BGB:AGB ratio. The BGB:AGB ratio increased with tree size largely due to an increase in small roots.The proportion of total root mass in the small roots (<15 mm) obtained through coring was 25–48% with 18–30% of total root biomass in the <5 mm diameter class.     

Influence of individual reserve trees on nearby reproduction in two-aged Appalachian hardwood stands

In the 1970s, public opposition to clearcut harvesting in hardwood forests of the eastern United States led forest managers and scientists to consider alternative practices that retain a low-density overstory forest cover. From 1979 to 1984, a form of clearcut-with-reserves harvesting was applied in 80-year-old Appalachian mixed-hardwoods to create four experimental stands with two-aged structures. The residual stand basal area averaged 5.3 m²/ha, comprising an average of 36 reserve trees/ha. The reserve trees were evenly distributed throughout the stand, initially with considerable space between their crowns, thus providing the sunlight and seedbed conditions needed to recruit desirable shade-intolerant reproduction after harvest. This study examined the response of the 100-year-old reserve trees and the development of the 20-year-old natural reproduction located in their immediate vicinity.Diameter at breast height (Dbh), height, and relative position were recorded for all reproduction ≥2.5 cm within transects adjacent to northern red oak (Quercus rubra L.) and yellow-poplar (Liriodendron tulipifera L.) reserve trees. Each transect was divided into five zones, which represented positions relative to the reserve tree crown edge, and basal area was computed for each of three shade tolerance classes within each zone. A repeated measures ANOVA was used to compare basal area of reproduction by tolerance classes and zone. In general, basal area of reproduction, particularly that of shade-intolerant species, increased with distance from the reserve tree. Regression analyses also indicated that dbh and height of reproduction was positively related to distance from the reserve trees. Although height growth of reserve trees was similar for both species, northern red oak exhibited significantly greater dbh and crown radial growth than yellow-poplar.The results indicated that reserve trees influence the growth rate and species composition of reproduction in their immediate vicinity. Basal area of reproduction increased from 10.1 to 17.7 m²/ha with increasing distance from the reserve trees. Basal area of intolerant species more than doubled along the same gradient. Basal area of reproduction in the two-age stands was 30–40% less than that observed in even-aged stands on similar growing sites, but the reduction was offset by growth of the reserve trees. The surface area covered by the reserve tree crowns increased approximately 88% for northern red oak and 44% for yellow-poplar. Since the sphere of influence of reserve trees increases over time, forest managers must consider their long-term impact on reproduction when prescribing clearcut-with-reserves harvests and other practices that involve retaining trees for many years.     

Stand and landscape level effects of a major outbreak of spruce beetles on forest vegetation in the Copper River Basin,Alaska

From 1989 to 2003, a widespread outbreak of spruce beetles (Dendroctonus rufipennis) in the Copper River Basin, Alaska, infested over 275,000 ha of forests in the region. During 1997 and 1998, we measured forest vegetation structure and composition on one hundred and thirty-six 20-m × 20-m plots to assess both the immediate stand and landscape level effects of the spruce beetle infestation. A photo-interpreted vegetation and infestation map was produced using color-infrared aerial photography at a scale of 1:40,000. We used linear regression to quantify the effects of the outbreak on forest structure and composition. White spruce (Picea glauca) canopy cover and basal area of medium-to-large trees [≥15 cm diameter-at-breast height (1.3 m, dbh)] were reduced linearly as the number of trees attacked by spruce beetles increased. Black spruce (Picea mariana) and small diameter white spruce (<15 cm dbh) were infrequently attacked and killed by spruce beetles. This selective attack of mature white spruce reduced structural complexity of stands to earlier stages of succession and caused mixed tree species stands to lose their white spruce and become more homogeneous in overstory composition. Using the resulting regressions, we developed a transition matrix to describe changes in vegetation types under varying levels of spruce beetle infestations, and applied the model to the vegetation map. Prior to the outbreak, our study area was composed primarily of stands of mixed white and black spruce (29% of area) and pure white spruce (25%). However, the selective attack on white spruce caused many of these stands to transition to black spruce dominated stands (73% increase in area) or shrublands (26% increase in area). The post-infestation landscape was thereby composed of more even distributions of shrubland and white, black, and mixed spruce communities (17–22% of study area). Changes in the cover and composition of understory vegetation were less evident in this study. However, stands with the highest mortality due to spruce beetles had the lowest densities of white spruce seedlings suggesting a longer forest regeneration time without an increase in seedling germination, growth, or survival.     

Dead trees and protected polypores in unmanaged north-temperate forest stands of Lithuania

The availability of coarse woody debris (CWD) and distribution of dead trees into categories of mortality (dead standing, broken and uprooted) were investigated in north-temperate forests of central Europe (Lithuania). The studied area comprised 188.7 ha and included 18 different stands 40–130 years of age with a variety of tree species (spruce (Picea abies (L.) Karst.), pine (Pinus sylvestris L.), alder (Alnus glutinosa (L.) Gaertn.), birch (Betula pendula Roth and B. pubescens Ehrh.), aspen (Populus tremula L.), oak (Quercus robur L.), forest types (caricus-sphagnum, vaccinium-myrtillus, oxalis, myrtillus-oxalis, caricus-calamagrostis) and edaphic conditions (peaty, sandy, loamy soils of different moisture). The stands were excluded from wood harvesting for at least 30 years. A total of 11 365 dead trees (over 10 cm in DBH) or 6160.7 m³ of dead wood was found (60.2 trees/ha and 32.6 m³/ha). The volume of CWD per hectare was larger in older stands (r_S=0.78, P<0.01). Tree mortality during the last 2 years consisted of 482 trees and 381 m³, or 1.28 trees/ha×year and 1.01 m³/ha×year. In 25–33% of cases it was wind-related. Uprooted and broken trees were of larger DBH than dead standing. The distribution into the categories of mortality was strongly dependent on tree species (chi-square test, d.f.=10,P=0). Dead standing dominated in CWD of pine and alder. Broken trees comprised almost a half in CWD of aspen, and about one-third in birch, alder and oak. Uprooting most often occurred in spruce, aspen and birch. Edaphic conditions and stand age had a pronounced impact on distribution into mortality categories for spruce (chi-square test, d.f.=20, P<0.00001) and pine (d.f.=8, P≤0.0003). On peat soil, only a minority of trees of both pine and spruce was uprooted, and standing dead prevailed. In CWD of spruce and pine, the proportions of both dead standing and broken decreased and that of uprooted trees increased on mineral soils of higher moisture and bulk density in older stands. By contrast, uprooting in birch and alder occurred less often on more wet sites, where the proportions of standing snags were higher. A total of 41 species of wood-decomposing polypores were found in the study area. Among those, 10 (24%) were of conservation value.     

Can intensive management accelerate the restoration of Brazil's Atlantic forests?

Only 7% of the once extensive forest along the eastern coast of Brazil remains, and much of that is degraded and threatened by agricultural expansion and urbanization. We wondered if methods similar to those developed to establish fast-growing Eucalyptus plantations might also work to enhance survival and growth of rainforest species on degraded pastures composed of highly competitive C₄ grasses. An 8-factor experiment was laid out to contrast the value of different intensities of cultivation, application of fertilizer and weed control on the growth and survival of a mixture of 20 rainforest species planted at two densities: 3 m × 1 m, and 3 m × 2 m. Intensive management increased seedling survival from 90% to 98%, stemwood production and leaf area index (LAI) by ～4-fold, and stemwood production per unit of light absorbed by 30%. Annual growth in stem biomass was closely related to LAI alone (r² = 0.93, p < 0.0001), and the regression improved further in combination with canopy nitrogen content (r² = 0.99, p < 0.0001). Intensive management resulted in a nearly closed forest canopy in less than 4 years, and offers a practical means to establish functional forests on abandoned agricultural land.     

A non-invasive method for reconstructing the relative mortality rates of trees in mixed-age,mixed-species forests

The death of overstory trees drives changes throughout forest ecosystems. Knowledge of mortality rates for these larger trees provides a long-term perspective on forest development and forest health. Tree mortality rates are typically determined by repeated censusing of trees over extended time intervals. We describe a method of reconstructing relative mortality rates that does not require injuring live trees on the study plots. To determine when trees died, we used cross-dating (matching tree-ring patterns of dead and live trees), identification of sapling releases, and assessment of tree decomposition. Dead trees were identified to genus or sub-genus by microscopic examination of wood. We reconstructed live tree community structure so that mortality could be relativized on a taxonomic-group- and tree-size-specific basis by the tree densities existing when mortality occurred. We modified a forest development model to operate backwards in time and validated this model by comparing past tree stem diameters predicted by the model with past stem diameters known from tree-ring measurements. All tree-ring measurements from live trees were obtained off the study plots. We report data for all trees ≥20 cm diameter at breast height (DBH) from seven oak-hickory forest sites in Arkansas, Illinois, Indiana, and Ohio, USA. Total plot area was 6.7 ha. Mortality rates were determined for the past 20 years by genus or sub-genus, DBH, and decade. The average mortality rate was 0.60% per year. The accuracy of the reconstructed mortality rates approaches that of mortality rates obtained by repeated censusing. While this mortality reconstruction method does not yield data for individual species, and may involve some compromise in accuracy, the method offers substantial benefits: long-term mortality data may be obtained from a short-term study, mortality rates may be obtained for the past, and because this reconstruction method is non-invasive, future mortality rates can be measured on the same plots.     

Diameter increment patterns among 106 tree species in a logged and silviculturally treated Costa Rican rain forest

Studies of growth rates of trees in managed neotropical forests have rarely employed complete botanical identification of all species, while published information for Central American lowland rain forests largely concerns forests free of recent disturbance. We studied diameter increments of trees in a managed Costa Rican rain forest. The Pentaclethra macroloba-dominated forest was located on low hills with Ultisols in Holdridge's Tropical Wet Forest life zone. The 540 m × 540 m (29.2 ha) experimental area was lightly logged during 1989–1990. The 180 m × 180 m (3.24 ha) experimental plots comprised a 100 m × 100 m (1.0 ha) central permanent sample plot (PSP) with a 40-m wide buffer strip. Post-harvest silvicultural treatments were liberation/refinement (in 1991) and shelterwood (in 1992), applied under a complete randomized block design with three replicates, using logged but untreated plots as controls. All live trees ≥10 cm DBH in the PSPs, were identified to species; data reported are for 1993–1996. Cluster analysis was used to group species on the basis of the median and quartiles of their diameter increment distributions, separating data by silvicultural treatments; five diameter increment groups were established and subdivided on the basis of the adult height of each species (four categories), giving 17 species groups in the final classification. Adult height and silvicultural treatment made a significant contribution to growth rate variation. Median annual increments of the slowest-growing species groups, which featured many under- and middle story species, were ca. 1 mm; those for the fastest growing species, which were mainly canopy and emergents, were ca. 16 mm. All species in the groups of very fast growth were pioneers, whether short or long-lived, though many other pioneer species did not show fast growth. The proportions of species found in groups of moderate, fast or very fast growth were greater in the silviculturally treated plots than in the controls, and one complete diameter increment group, of fast growth, was only represented in the treated plots. Crown form, crown illumination and presence of lianas in the crown, showed significant correlations with diameter increments, though the importance of these latter two variables varied with silvicultural treatment. The very fast growth groups differed from the others in having higher proportions of trees with well-formed, well-illuminated crowns and an irregular diameter distribution with relatively few individuals in the smallest DBH class. Comparison with data from other neotropical forest sites shows that long-lived pioneers such as Vochysia ferruginea and Jacaranda copaia grow fast or very fast at all sites, while non-commercial canopy and emergent species of Chrysobalanaceae and Sapotaceae appear to be uniformly slow-growing. Growth data for the majority of species are, however, published for the first time.     

First decadal response to treatment in a disturbance-based silviculture experiment in Maine

Disturbance-based silvicultural systems generally seek to promote complex stand structures that are consistent with temporal and spatial patterns of natural disturbance while allowing for the sustainable harvest of timber. Gap-based harvesting systems are commonly used within this framework because they can be designed to approximate the frequencies and spatial patterns of a wide array of disturbance regimes. Patterns in stand-level growth, sapling recruitment and regeneration response were examined for one such gap-based system, the Acadian Forest Ecosystem Research Program (AFERP) in central Maine, that was designed to emulate the annual 1% disturbance frequency typical of the northeastern United States and Canada. A decade after treatment, stand-level differences in basal area growth and density between two gap-based treatments and an unharvested control were not statistically significant, largely due to low replication, but within-stand growth and regeneration responses differed strongly by spatial position relative to harvest gaps. Regeneration of shade-tolerant and intolerant species increased regardless of gap size, likely a response to increasing light availability from canopy openings due to harvesting and mortality. Further, there was evidence of gap size effects on sapling recruitment as large gaps (>1000 m²) favored the growth and survival of mid-successional species such as red maple and white pine, while small gaps (<1000 m²) favored late-succession species such as eastern hemlock and spruce. Overstory growth rates also differed by both species and position relative to harvest gaps with most species growing best in gaps and better along gap edges than in adjacent forest. Notably, overstory growth rates for white pine were not influenced by spatial position. These results suggest harvest gaps may have significant growth and regeneration impacts in adjacent, yet unharvested areas, which could lead to profound differences in forest development over the rotation. Obviously, longer-term studies of gap-based systems are needed to more clearly elucidate these responses.     

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.