Natural development and regeneration of a Central European montane spruce forest

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

Topographic and compositional influences on disturbance patterns in a northern Maine old-growth landscape

Disturbance patterns are strongly coupled with forest composition and structure, and patterns change through time in response to shifts in climate, anthropogenic impacts and other factors. Knowledge of the natural disturbance patterns for establishing baseline conditions for a forest type or ecosystem facilitates change detection for other elements of the biophysical system important to management and conservation. Dendrochronological reconstructions from old-growth forest remnants throughout northeastern North America document average decadal rates of disturbance of 5%–<10% over the last 150–300 years. Relatively frequent, low severity disturbance characterized by small gaps representing canopy openings made by 1–3 trees prevail in these forests dominated by varying mixtures of late-successional tree species. Few studies, though, have explicitly characterized differences related to composition or topographic setting in old-growth landscapes. We addressed this by comparing the temporal and spatial disturbance patterns reconstructed from tree rings at two spatial resolutions (0.5 ha and 200 m²). Sites were selected to assess the influence of topography (slope) and cover type in stands where red spruce (Picea rubens Sarg) and balsam fir (Abies balsamea L. Mill) were key components. Low rates of disturbance (average <10% per decade) and small gap sizes (≤30 m²) prevailed in all stands during the decades from 1850–1980. Episodic pulses of disturbance, of nearly moderate intensity in some stands, opened ca. 20–30% of the canopy area and were associated with wind events and/or insect outbreaks that differentially affected stands. We found no significant difference in the average temporal rates of disturbance related to cover type or topography in 0.5-ha plots. However, the influence of these factors was evident in comparisons of gap areas estimated for 200-m² plot sections. At this resolution, the largest canopy openings (≥100 m²) occurred most frequently in slope sites, enabling pulses of canopy accession for Betula alleghaniensis (Britton). Whereas the smallest canopy openings (≤30 m²) dominated softwood stands, favoring red spruce, balsam fir and other shade-tolerant species throughout the forest. The variable effects of common disturbance agents, regardless of topographic position and/or cover types, points to the important role of biological legacies determining stand structure and composition on subsequent disturbance events and long-term patterns.     

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.     

A simulation of the development and restoration of old-growth structural features in northern hardwoods

Thinning treatments in second-growth forest may be a practical means of accelerating the development of certain old-growth structural features in regions where old stands are presently uncommon. We used CANOPY, an individual-tree model calibrated with data from thinned and unthinned stands, to simulate effects of thinning on growth rates and development of old-growth structural features in second-growth northern hardwoods. Three simulated, moderately heavy thinnings over a period of 45 years nearly doubled the predicted mean radial increment of canopy trees, percent of stand basal area in large trees, and area of canopy gaps. Compared to untreated stands, thinned stands had fewer dead trees per ha, but the dead trees were larger in size and the overall volume of snags and logs was little affected. In a 77-year old even-aged stand, moderately heavy thinning was predicted to reduce the time needed to attain the minimum structural features of an old-growth forest from 79 to 36 years. Simulated treatments in an older, uneven-aged stand gave mixed results; the moderately heavy treatment stimulated individual tree growth, but the removal of some medium-sized canopy trees in conjunction with natural mortality delayed the development of old-growth structure. Total volume of dead wood may still be deficient under the thinning regimes investigated in this study, but predicted live-tree structure 45 years after moderately heavy thinning was typical of stands in the advanced transition and steady-state stages of old-growth development. Results suggest that thinning can substantially accelerate the development of old-growth structure in pole and mature northern hardwoods, but response in older, uneven-aged stands is more modest, and treatments in these stands may need to be more conservative to achieve restoration goals.     

The role of gaps and tree regeneration in the transition from dense to open black spruce stands

Black spruce forests growing on clay soils in northwestern Quebec change structure from dense even-aged stands to open uneven-aged stands such that almost all forests older than 200 years have an open canopy. These forests become unproductive over time because they are prone to paludification. The main goal of our study was to document the transition between dense and open stands in terms of gap dynamics, with a focus on tree regeneration. Our objective was to determine whether forests remain open due to a lack of regeneration, a lack of growth or both. Nine stands along a 50–250-year-old time since fire gradient were sampled with the line intersect sampling method. Gap fraction increased with stand age and reached a maximum of 77% in the oldest site. In old-growth stands, gaps were interconnected due to the low density of these forests. Most of the gap makers were found with broken stems. Regeneration was dominated by black spruce layers and was relatively abundant (1.71 stems/m²). However, the majority of gap fillers were smaller than 1 m in height in stands of all ages. Instead of a lack of regeneration, the opening of the forests is due to a lack of growth associated with cold and wet organic deposits. Partial harvesting could be implemented on the most productive sites, while management techniques including soil disturbances will be required on low productivity sites to recreate good growth conditions.     

The ecology of tree regeneration in mature and old forests: combined knowledge for sustainable forest management

We focused our attention on quantifying the factor complex of forest regeneration in 423 mature and old stands with contrasting environmental conditions. We recorded the microhabitat selection of tree recruits, the frequency of tree seedlings, and evaluated the drivers of sapling abundance and diversity. The majority of forest regeneration was established on undisturbed forest floor. Dead wood was a frequent substrate in spruce-(co)dominated forests. Seedling frequency within a stand was related to the site-type specific productivity gradient of stands—pine seedlings were common in low-productivity and spruce in high-productivity boreal forests. Seedlings of temperate broad-leaved trees dominated in productive boreonemoral forests, except for oak, which showed a uniform distribution of abundance in all forest site-types. Sapling abundance was dictated by forest site-type, and facilitated by stand diversity, variability in stand closure, lying dead wood, abundant moss, and a thick organic layer. Only in boreal forests was sapling abundance suppressed by the abundant spruce and younger trees. Upon considering the relationship between sapling abundance and species richness, sapling diversity was dependent on forest site-type, suppressed by stand density and dead wood (old gap) abundance, and facilitated by stand diversity. In addition, boreonemoral stands, competition from herbs, and facilitation by mosses occurred. The observed pattern of tree recruitment points to the importance of top-down effects of the overstory, competing or facilitating interactions with forest floor vegetation, and availability of regeneration microhabitats, which in complex make their ecology comparable with forest herbs. Natural forest regeneration can be enhanced if silvicultural methods support mixed stands and enhance field layer diversity. Oak can provide the universal tree species to improve stand structure over a wide range of habitats.     

Comparing height growth and biomass production of black spruce trees in logged and burned stands

The strategy of using advanced layering for regenerating logged black spruce stands has become a common practice. Compared with natural post-fire black spruce stands, this strategy may alter stand structure with a possible change in stand productivity. Using harvested tree data from sample plots established in burned and logged stands, 50 years after disturbance, and on similar soils, we compared the structure, height growth, and biomass allocation of both types of stands. Stem analysis revealed that black spruce trees in logged stands reached their maximum height growth later and at a concomitant lower level than black spruce trees in burned stands. Biomass production was comparable between stand types but was reduced when ericaceous shrubs were abundant. Compared to natural post-fire stands, logged stands present a shift of biomass allocation to branches and to leaves. These mechanisms are interdependent and represent the tree adjustment to the altered structure in logged stands characterised by the fragmented canopy with the increase of ericaceous shrubs cover and of organic layer thickness.     

Reconstructing 100-150 years of logging history in coastal spruce forest (Picea abies) with special conservation values in central Norway

Coastal spruce forests of central Norway harbour a unique assemblage of epiphytic lichens and are given high priority with respect to conservation of biodiversity. To assess the historical impact of logging during the last 100-150 yrs, 31 remnant stands were studied by means of tree-ring analysis of 2199 trees and the decay stage of 1605 stumps. No stands had been clear-cut, but all had been selectively logged at least twice during the last 150 yrs. Total harvested timber volume ranged from 65 to 409 m³ha-¹ (31-124% of present-day standing volume) and the selective logging kept standing volume low (40-200 m³ha-¹) during 1890-1930. Present-day stand characteristics were strongly correlated with site productivity and topographic position within the ravine valleys. Low amounts of dead wood at sites with high historical logging activity was the only consistent relationship found after covariance of site productivity, topographic position and deciduous trees were taken into account. The results indicate that old-growth stand characteristics, such as reversed J-shaped age distributions and dead wood in advanced decay classes, can be obtained 100-150 yrs after intensive selective logging.     

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.     

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.     

Storm legacies shaping post-windthrow forest regeneration: learnings from spatial indices in unmanaged Norway spruce stands

The anticipated increase in extreme disturbance events due to climate change is likely to expose Norway spruce (Picea abies (L.) Karst.) dominated forests in northern Europe to new conditions. Empirical data on the resilience of such natural (unmanaged) forests to disturbance and the long-term patterns of regeneration in its aftermath are currently scarce. We performed a quantitative assessment of natural forest stands in north–western Latvia to identify and characterise the patterns of stand structure 44 years after a stand-replacing disturbance and investigated the effects of legacies on regeneration. The spatial distribution of tree species and their dimensions were assessed in 71 circular sample plots (500 m² each) in natural forest areas, where Norway spruce dominated prior to the windthrow and salvage logging was not carried out. Spatial indices (species mingling index, size differentiation index, and aggregation index) were used to characterise stand structure and diversity. The different initial states (age and coverage of surviving trees) of stands affected eventual tree species dominance, size differentiation, degree of mingling and aggregation. Our results demonstrate a close relationship between disturbance legacies and spatial indices. The pre-storm understory and canopy survivors decreased species mingling, whereas survivors increased size differentiation. The size differentiation increased also with a higher degree of species mingling. Leaving differential post-storm legacies untouched promotes a higher structural and species diversity and therefore supports the management approach of preserving canopy survivors.

     

The effects of partial cutting on stand structure and growth of western hemlock–Sitka spruce stands in southeast Alaska

The effects of partial cutting on species composition, new and residual-tree cohorts, tree size distribution, and tree growth was evaluated on 73 plots in 18 stands throughout southeast Alaska. These partially cut stands were harvested 12–96 years ago, when 16–96% of the former stand basal area was removed.Partial cutting maintained stand structures similar to uncut old-growth stands, and the cutting had no significant effects on tree species composition. The establishment of new-tree cohorts was positively related to the proportion of basal-area cut. The current stand basal area, tree species composition, and stand growth were significantly related to trees left after harvest (p<0.001). Trees that were 20–80 cm dbh at the time of cutting had the greatest tree-diameter and basal-area growth and contributed the most to stand growth. Diameter growth of Sitka spruce and western hemlock was similar, and the proportion of stand basal-area growth between species was consistent for different cutting intensities.Concerns about changing tree species composition, lack of spruce regeneration, and greatly reduced stand growth and vigor with partial cuts were largely unsubstantiated. Silvicultural systems based on partial cutting can provide rapidly growing trees for timber production while maintaining complex stand structures with mixtures of spruce and hemlock trees similar to old-growth stands.     

Long-term dynamics of a mixed conifer stand in Slovenia managed with a farmer selection system

The single-tree selection system is an important option for management of Norway spruce (Picea abies (L.) Karst.) and silver fir (Abies alba Mill.) forests because it provides continuous cover, requires low investments for tending, and promotes natural regeneration as well as high stand resistance and elasticity. It is often regarded as a very conservative system that usually results in only minor spatiotemporal changes in forest structure and composition. We studied management history, structural changes, regeneration dynamics, and light climate of a traditional single-tree farmer selection silver fir-Norway spruce forest (site typology Bazzanio-Abietetum). Stand structure was analyzed on five 0.25 ha permanent plots in 1994, 2001, and 2008. Regeneration density and height growth, forest floor vegetation, and light climate were also assessed on 1.5 × 1.5 m regeneration subplots in 2001 and 2008. Tree cores extracted from dominant trees from both species in two plots were used for reconstructing stand history and age structure of the canopy layer. We documented the forest response to three types of selection management regimes: excessive, normal, and conservative. Excessive management with harvest intensity significantly above the increment was documented until the late 1950s, including two peaks of heavy fellings (diameter limit cut) in the 1880s and 1930s, which favoured establishment of Norway spruce and released regeneration. The period that followed was characterized by normal selection management, but was nevertheless marked by a decline of silver fir as a result of air pollution and several droughts. This led to sanitary fellings that were carried out from the late 1970s to the early 1990s. In the last two decades conservative management followed, which led to suppression and decline of regeneration, especially of Norway spruce, and loss of selection structure. Although we recorded lower regeneration potential of silver fir compared with Norway spruce within the seedling category, silver fir outcompeted Norway spruce within the small-sized tree category (1 cm < dbh ? 10 cm) because of its superior height growth in low light levels (diffuse light <6%) and occupied a greater share of the canopy. Nevertheless, we anticipate that over the long-term the low light regime will also cause regeneration decline of silver fir and broadleaves. Our research revealed significant structural changes in a single-tree farmer selection forest during the last 150 years. These were a result of variable management regime and environment. A farmer single-tree selection system could better mimic the natural disturbance regime if spatiotemporal combinations of diverse felling regimes would be used.     

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.     

Photosynthetic characteristics of Fagus sylvatica and Quercus robur established for stand conversion from Picea abies

Efforts in Europe to convert Norway spruce (Picea abies) plantations to broadleaf or mixed broadleaf-conifer forests could be bolstered by an increased understanding of how artificial regeneration acclimates and functions under a range of Norway spruce stand conditions. We studied foliage characteristics and leaf-level photosynthesis on 7-year-old European beech (Fagus sylvatica) and pedunculate oak (Quercus robur) regeneration established in open patches and shelterwoods of a partially harvested Norway spruce plantation in southwestern Sweden. Both species exhibited morphological plasticity at the leaf level by developing leaf blades in patches with an average mass per unit area (LMA) 54% greater than of those in shelterwoods, and at the plant level by maintaining a leaf area ratio (LAR) in shelterwoods that was 78% greater than in patches. However, we observed interspecific differences in photosynthetic capacity relative to spruce canopy openness. Photosynthetic capacity (A₁₆₀₀, net photosynthesis at a photosynthetic photon flux density of 1600 μmol photons m⁻² s⁻¹) of beech in respect to the canopy gradient was best related to leaf mass, and declined substantially with increasing canopy openness primarily because leaf nitrogen (N) in this species decreased about 0.9 mg g⁻¹ with each 10% rise in canopy openness. In contrast, A₁₆₀₀ of oak showed a weak response to mass-based N, and furthermore the percentage of N remained constant in oak leaf tissues across the canopy gradient. Therefore, oak photosynthetic capacity along the canopy gradient was best related to leaf area, and increased as the spruce canopy thinned primarily because LMA rose 8.6 g m⁻² for each 10% increase in canopy openness. These findings support the premise that spruce stand structure regulates photosynthetic capacity of beech through processes that determine N status of this species; leaf N (mass basis) was greatest under relatively closed spruce canopies where leaves apparently acclimate by enhancing light harvesting mechanisms. Spruce stand structure regulates photosynthetic capacity of oak through processes that control LMA; LMA was greatest under open spruce canopies of high light availability where leaves apparently acclimate by enhancing CO₂ fixation mechanisms.     

Semi-forest coffee cultivation and the conservation of Ethiopian Afromontane rainforest fragments

Coffea arabica shrubs are indigenous to the understorey of the moist evergreen montane rainforest of Ethiopia. Semi-forest coffee is harvested from semi-wild plants in forest fragments where farmers thin the upper canopy and annually slash the undergrowth. This traditional method of coffee cultivation is a driver for preservation of indigenous forest cover, differing from other forms of agriculture and land use which tend to reduce forest cover. Because coffee farmers are primarily interested in optimizing coffee productivity, understanding how coffee yield is maximized is necessary to evaluate how, and to what extent, coffee production can be compatible with forest conservation.Abiotic variables and biotic variables of the canopy were recorded in 26 plots within 20 forest fragments managed as semi-forest coffee systems near Jimma, SW Ethiopia. In each plot, coffee shrub characteristics and coffee yield were recorded for four coffee shrubs. Cluster and indicator species analyses were used to differentiate plant communities of shade trees. A multilevel linear mixed model approach was then used to evaluate the effect of abiotic soil variables, shade tree plant community, canopy and stand variables, coffee density and coffee shrub size variables on coffee yield.Climax species of the rainforest were underrepresented in the canopy. There were three impoverished shade tree communities, which differed in tree species composition but did not exhibit significant differences in abiotic soil variables, and did not directly influence coffee yield. Coffee yield was primarily determined by coffee shrub branchiness and basal diameter. At the stand level a reduced crown closure increased coffee yield. Yield was highest for coffee shrubs in stands with crown closure less than median (49 ± 1%). All stands showed a reduced number of stems and a lower canopy compared to values reported for undisturbed moist evergreen montane rainforests.Traditional coffee cultivation is associated to low tree species diversity and simplified forest structure: few stems, low canopy height and low crown closure. Despite intensive human interference some of the climax species are still present and may escape local extinction if they are tolerated and allowed to regenerate. The restoration of healthy populations of climax species is critical to preserve the biodiversity, regeneration capacity, vitality and ecosystem functions of the Ethiopian coffee forests.     

Growth and productivity of black spruce in even- and uneven-aged stands at the limit of the closed boreal forest

The increasing commercial interest and advancing exploitation of new remote territories of the boreal forest require deeper knowledge of the productivity of these ecosystems. Canadian boreal forests are commonly assumed to be evenly aged, but recent studies show that frequent small-scale disturbances can lead to uneven-aged class distributions. However, how age distribution affects tree growth and stand productivity at high latitudes remains an unanswered question. Dynamics of tree growth in even- and uneven-aged stands at the limit of the closed black spruce (Picea mariana) forest in Quebec (Canada) were assessed on 18 plots with ages ranging from 77 to 340 years. Height, diameter and age of all trees were measured. Stem analysis was performed on the 10 dominant trees of each plot by measuring tree-ring widths on discs collected each meter from the stem, and the growth dynamics in height, diameter and volume were estimated according to tree age. Although growth followed a sigmoid pattern with similar shapes and asymptotes in even- and uneven-aged stands, trees in the latter showed curves more flattened and with increases delayed in time. Growth rates in even-aged plots were at least twice those of uneven-aged plots. The vigorous growth rates occurred earlier in trees of even-aged plots with a culmination of the mean annual increment in height, diameter and volume estimated at 40–80 years, 90–110 years earlier than in uneven-aged plots. Stand volume ranged between 30 and 238 m³ ha⁻¹ with 75% of stands showing values lower than 120 m³ ha⁻¹ and higher volumes occurring at greater dominant heights and stand densities. Results demonstrated the different growth dynamics of black spruce in single- and multi-cohort stands and suggested the need for information on the stand structure when estimating the effective or potential growth performance for forest management of this species.     

Canopy dynamics in unlogged and logged forest stands in the eastern Amazon

Selective logging of tropical forests damages residual trees and creates canopy openings throughout the stand. In the Amazon, reduced-impact logging results in significantly less damage than conventional unplanned logging; yet either logging method leads to substantial fragmentation of the forest canopy. Increased mortality rates of trees damaged in logging have been documented. In this paper, we investigate the effect of logging disturbance on treefall rates.Using repeat inventories of canopy trees (≥35 cm dbh) in six large (≥50-ha) forest stands at two sites in the eastern Amazon, we measured mortality rates in three treatment classes: unlogged, conventionally logged (CL) and logged using reduced-impact methods (RIL). At least 3000 trees were mapped and inventoried per stand prior to timber harvests. In the second inventory, 3 years after harvests in the logged stands, all trees were located and scored as living, standing dead, uprooted or broken.We found significantly higher overall mortality rates for trees in logged forest (both CL and RIL stands) than in adjacent unlogged forest. This effect was largely due to higher rates of treefall (i.e., stems uprooted or broken from natural causes). Moreover, significantly higher treefall rates were recorded for trees in logged forest that were not damaged in the logging than for trees in unlogged stands. Treefall rates were nearly twice as high in conventionally logged forest as in forest logged using reduced-impact methods. We found indirect support for the hypothesis that increased treefall rates in logged forest are related to increased canopy opening and fragmentation through analysis of the locations of fallen trees in relation to canopy disturbance.     

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.