Nitrogen cycling following mountain pine beetle disturbance in lodgepole pine forests of Greater Yellowstone

Widespread bark beetle outbreaks are currently affecting multiple conifer forest types throughout western North America, yet many ecosystem-level consequences of this disturbance are poorly understood. We quantified the effect of mountain pine beetle (Dendroctonus ponderosae) outbreak on nitrogen (N) cycling through litter, soil, and vegetation in lodgepole pine (Pinus contorta var. latifolia) forests of the Greater Yellowstone Ecosystem (WY, USA) across a 0-30 year chronosequence of time-since-beetle disturbance. Recent (1-4 years) bark beetle disturbance increased total litter depth and N concentration in needle litter relative to undisturbed stands, and soils in recently disturbed stands were cooler with greater rates of net N mineralization and nitrification than undisturbed sites. Thirty years after beetle outbreak, needle litter N concentration remained elevated; however total litter N concentration, total litter mass, and soil N pools and fluxes were not different from undisturbed stands. Canopy N pool size declined 58% in recent outbreaks, and remained 48% lower than undisturbed in 30-year old outbreaks. Foliar N concentrations in unattacked lodgepole pine trees and an understory sedge were positively correlated with net N mineralization in soils across the chronosequence. Bark beetle disturbance altered N cycling through the litter, soil, and vegetation of lodgepole pine forests, but changes in soil N cycling were less severe than those observed following stand replacing fire. Several lines of evidence suggest the potential for N leaching is low following bark beetle disturbance in lodgepole pine.     

Long-term influence of fire and harvesting on boreal forest age structure and forest composition in eastern Québec

In boreal forests, historical variations in the area disturbed by natural disturbances or harvesting have rarely been compared. We measured temporal and spatial variations in areas affected by severe fires and clearcutting throughout the 20th century in a 57, 332 km² section of the eastern Canadian boreal forest. We examined the effects of these disturbances on spatio-temporal variations in the abundance of forests >60 years. Natural variability for the abundance of forests >60 years was estimated from simulations of natural disturbance regimes. We also measured compositional and structural differences between three categories of stands originating from relatively recent disturbances (∼50 years; clearcutting, fires, and clearcutting followed by fires), and one category of stands that were undisturbed for at least 200 years. At the regional level, we observed that forests >60 years gradually became scarcer throughout the 20th century due to a gradual expansion of harvested areas, an effect most pronounced in the southern part of the region, where mature and old forest abundance was clearly outside the range of natural variability at the end of the studied period. At the stand level, forest composition and structure differed between stand-origin categories: clearcutting-origin stands contained more balsam fir (Abies balsamea), fire-origin stands more black spruce (Picea mariana), and fire/clearcutting-origin stands more hardwoods (Betula papyrifera and Populus tremuloides). Overall, we estimate that strict forest management targets based on natural disturbance regimes will be difficult to achieve in eastern North-American boreal forests, most notably because contemporary disturbance rates, including both clearcutting and fire, have gradually become higher than the fire rates observed during the preindustrial period.     

Influence of fire and mountain pine beetle on the dynamics of lodgepole pine stands in British Columbia,Canada

An outbreak of the mountain pine beetle (Dendroctonus ponderosae Hopkins; MPB), currently affecting over 10.1 million hectares of lodgepole pine forests (Pinus contorta Dougl.) in British Columbia, Canada, is the largest in recorded history. We examined the dynamics of even-aged lodgepole pine forests in southern British Columbia, which were undergoing this MPB outbreak. Using dendroecology and forest measurements we reconstructed the stand processes of stand initiation, stand disturbances, tree mortality, and regeneration, and explained the current stand structure and the potential MPB impacts in selected stands. Our results indicate that stand-replacing fires initiated even-aged seral lodgepole pine stands in this region. In the absence of fire in the 20th century, multiple MPB disturbances, which each resulted in partial canopy removal, modified the simple one-layer structure of the fire-origin stands by the initiation of post-MPB disturbance regeneration layers, transforming the stands into complex, multi-aged stands. Despite high overstory mortality due to the current MPB outbreak, regeneration layers, which are likely to survive the current outbreak, will provide important ecological legacies and will contribute to mid-term timber supply.     

The composition and diversity of natural regeneration of tree species in gaps under different intensities of forest disturbance

The positive ecological interaction between gap formation and natural regeneration has been examined but little research has been carried out on the effects of gaps on natural regeneration in forests under different intensities of disturbance.This study evaluates the composition,diversity,regeneration density and abundance of natural regeneration of tree species in gaps in undisturbed,intermittently disturbed,and disturbed forest sites.Bia Tano Forest Reserve in Ghana was the study area and three gaps each were selected in the three forest site categories.Ten circular subsampling areas of 1 m2 were delineated at 2 m spacing along north,south,east,and west transects within individual gaps.Data on natural regeneration 350 cm height were gathered.The results show that the intensity of disturbance was disproportional to gap size.Species diversity differed significantly between undisturbed and disturbed sites and,also between intermittently disturbed and disturbed sites for Simpson's(1-D),Equitability(J),and Berger-Parker(B-P) indices.However,there was no significant difference among forest sites for Shannon diversity(H) and Margalef richness(MI) indices.Tree species composition on the sites differed.Regeneration density on the disturbed site was significantly higher than on the two other sites.Greater abundance and density of shade-dependent species on all sites identified them as opportunistic replacements of gap-dependent pioneers.Pioneer species giving way to shade tolerant species is a natural process,thus make them worst variant in gap regeneration.     

Natural regeneration characteristics of Pinus sylvestris var. mongolica forests on sandy land in Honghuaerji, China

Natural regeneration in Mongolian pine, Pinus sylvesttis var. mongolica, forest at Honghuaerji of China （the original of the natural Mongolian pine, forest on sandy land） was studied in 2004. The total mean values of regeneration indexes were higher in mature stands （more than 80% individual stems were older than 50 years）, the maximum of regeneration index reached 29 seedlings, m^ 2, with lowest values in the younger stand, e.g., in 32-year old and 43-year old stands. The stand age was an important factor determining the natural regeneration, which was the best in the older stands in this investigation （e.g. about 80-year old）. The regeneration index seemed not to be closely in relation to canopy openness although Mongolian pine is a photophilic tree species. In each type of gaps, natural regeneration was very well. Regeneration indexes were satisfactory at the south and east edges in the circle gaps; and at the east edge of the narrow-square gaps. Results indicated that Mongolian pine, seedlings could endure shading understory, but it would not enter the canopy layer without gap or large disturbance, e.g., fire, wind/snow damage or clear cutting etc. These results may provide potentially references to the management and afforestation of Mongolian pine, plantations on sandy land in arid and semi-arid areas. Researches such as the comprehensive comparisons on regeneration, structure and ecological conditions and so on between natural Mongolian pine, forests and plantations should be conducted in the future.     

Can a natural experiment be used as a tool to design partial cutting regimes? The decline of <Emphasis Type="Italic">Austrocedrus chilensis</Emphasis> forests,an example

The forests of Austrocedrus chilensis in Argentina suffer mortality from what is locally known as ‘mal del ciprés’. This disturbance usually appears as aggregations of dead trees of varying sizes. The mortality process seems unpredictable, and regeneration establishment is variable under these conditions. The highly variable mortality rates results in a natural experiment that allows the correlation of overstory mortality with regeneration rate. This correlation should then make it possible to estimate the regeneration success which would follow after different levels of partial cutting, allowing the purposeful direction of the regeneration process. We collected data from eight Austrocedrus chilensis-dominated stands and reconstructed overstory mortality over time and correlated it to the regeneration recruitment. Mortality showed two patterns: increasing ‘continuous’ mortality over time, and ‘pulses’ of important basal area loss in short periods. Overstory mortality had an effect on the recruitment and composition of the regeneration by allowing different species establishment depending on the degree of mortality. By quantifying mortality basal area and recruitment over time, we calculated estimates that can provide partial cutting guidance. Partial cuttings would then allow the purposeful direction of the regeneration process. Stand reconstruction studies provided a useful tool to help designing regeneration systems using a natural experiment. Some specific management implications for symptomatic A. chilensis forests are given.     

Spatial distribution,advanced regeneration and stand structure of Nepalese Sal (Shorea robusta) forests subject to disturbances of different intensities

We investigated the spatial distribution, advanced regeneration and stand structure of five Shorea robusta-dominated forests in 25 1-ha plots subject to disturbances of different intensities. We aim to elucidate the relationships of advanced regeneration and spatial patterns of the tree species with degree of disturbance magnitude. Sixty-seven tree species were recorded in the forest plots; 41 species were found in the least disturbed forests, while only 10 species were found in the heavily disturbed forests. We found 5320 trees with >1.5 cm diameter at breast height, in total, and found that moderately disturbed forests contained the highest advanced regeneration (sapling)/pole densities. No significant differences were observed in stem basal area among forests. The overall stand density changed quadratically across the disturbance gradient. A strong inverse relationship was found between the overall stand density and diameter class in the least disturbed and moderately disturbed forests. Ten species showed variation in their dispersion patterns across the disturbance gradient. Most of the socio-economically important tree species analyzed showed little or no regeneration in the least and most heavily disturbed forests. Individual species showed different responses to disturbance ranging from ‘tolerant’ (Shorea robusta, Lagerstroemia parviflora and Symplocos spp.) to ‘sensitive’ (Trewia nudiflora, Adina cardifolia and Terminalia alata). We concluded that moderate disturbance intensity not only ensures high stand density, but also enhances the advanced regeneration of socio-economically important tree species and affects their dispersion patterns. Future management strategy must balance the consumptive needs of the local community with those of species conservation by allowing regulated access to the forests.     

On the structural and species diversity effects of bark beetle disturbance in forests during initial and advanced early-seral stages at different scales

Following disturbances, early-seral stages of forests provide a variety of structures. Whether this variety is a short-term phenomenon or influences forest succession for several decades or even longer is not known. We tested the hypotheses that after spruce dieback caused by bark beetles, a high spatial heterogeneity of stand structures will persist within stands and among stands even in advanced early-seral stages and that species taxonomical and functional diversity measures will reflect this heterogeneity. We used a chronosequence of unmanaged forests in the Berchtesgaden National Park (Germany) consisting of mature undisturbed spruce stands (control), stands belonging to an initial early-seral stage (~3 years after disturbance) and stands in an advanced early-seral stage (~20 years after disturbance). We analysed diversity and heterogeneity of these forest stands including stand structure, species density, species composition and functional–phylogenetic diversity of vascular plants, wood-inhabiting fungi and saproxylic beetles within plots, among plots of the same successional stage and among stages. Stands of the advanced early-seral stage were characterized by a high spatial heterogeneity of structural attributes, such as crown cover, regeneration density and spatial distribution of trees. Among-plot taxonomic beta diversity was highest in the advanced early-seral stage for beetles, but lowest for fungi, while beta diversity of plants among plots remained unchanged during succession. The mosaic of successional stages initiated by bark beetles increased the gamma diversity of the study area, especially for fungi and beetles. Our findings support the hypothesis that structural heterogeneity continues for at least two decades at stand and landscape scales and that species turnover among successional stages is a major mechanism for gamma diversity in forests after bark beetle disturbance.     

Scarification and Seedfall affects Natural Regeneration of Scots Pine Under Two Shelterwood Densities and a Clear-cut in Southern Sweden

A study was undertaken to evaluate the effect of Scots pine (Pinus sylvestris) shelterwood density and timing of removal on the regeneration of forests with improved wood quality. This paper focuses on the effect of scarification and seedfall on the success of natural regeneration of Scots pine under two shelterwood densities and in a clear-cut. Wood quality aspects will be addressed later in the study. After 4 yrs, natural regeneration of Scots pine under a 200 stems ha^-1 shelterwood reached 90000 seedlings ha^-1, 53000 under a 160 stems ha^-1 shelterwood and 3700 in a clear-cut. The high natural regeneration under the densest shelterwood resulted from a high seedfall, slower invasion by competing vegetation, consequent prolonged recruitment and low mortality. Since mortality largely decreased for both shelterwood densities and recruitment seems to continue, the success of regeneration should be maintained in the future if proper release operations are conducted. The latter could begin when seedlings reach a height of about 0.5 m and the cover should be maintained until they reach a height of about 6 m. Therefore, it may be possible to use 200 stems ha-1 Scots pine shelterwoods in southern Sweden to obtain dense stands and thus improve wood quality (stem taper, wood density, etc.).     

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.     

Silvicultural systems for southern bottomland hardwood forests

Silvicultural systems integrate both regeneration and intermediate operations in an orderly process for managing forest stands. The clearcutting method of regeneration favors the development of species that are moderately intolerant to intolerant of shade. In fact, clearcutting is the most proven and widely used method of successfully regenerating bottomland oak species in the South. The seed-tree method of regeneration favors the establishment of light-seeded species. Mechanical soil scarification may be necessary if the desired species requires bare mineral soil for establishment. The shelterwood method of regeneration can provide for the development of heavy-seeded species, but has produced highly variable results with southern bottomland oaks. The single-tree selection method of regeneration favors the development of shade-tolerant species. When single-tree selection is applied repeatedly to stands containing commercially valuable shade-intolerant species, composition will gradually shift to less-valuable, more-tolerant species. Consequently, the single-tree selection method of regeneration is not recommended for any commercially valuable bottomland hardwood tree species. Group selection, in its strictest application, creates only small openings that usually fail to allow sufficient light to the forest floor for satisfactory establishment and development of shade-intolerant bottomland species. Patch cutting, a combination of uneven-aged (group selection) and even-aged (clearcutting) silviculture, designed to create larger openings, has been successfully used to produce an uneven-aged stand that consists of many small, irregularly shaped, even-aged groups. Silvicultural systems should include a planned program of intermediate operations designed to enhance the growth and development of those species favored during the regeneration process. Improvement cutting and commercial thinning are increasingly common in southern bottomland hardwood forests. Other partial cuttings employed today in bottomland hardwood forests typically involve some form of crop-tree release. Specific recommendations for the selection of silvicultural systems are presented for the eight most important species groups found in southern bottomland hardwood forests.     

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.     

Short term effects of alternative silvicultural treatments on stand attributes in irregular balsam fir-black spruce stands

The eastern Canadian boreal forest exhibits a specific disturbance regime where forest fires are less frequent than in the western part. This particularity may explain the abundance of irregular stands with distinct ecological features. To ensure sustainable forest management, these characteristics require the implementation of an adapted silviculture regime. In this context, two selection cutting methods were developed and compared with more conventional techniques, initially designed for cutting more regular stands of the boreal forest (cutting leaving small merchantable stems, careful logging preserving advance regeneration). The comparison focused on the capacity of treatments to maintain the primary attributes of irregular boreal forests, including complex vertical structure, abundant tree cover, species composition, and an abundance of dead wood. Mortality and regeneration processes were also compared.     

Spatial and temporal patterns in structure,regeneration, and mortality of an old-growth ponderosa pine forest in the Colorado Front Range

Effective management and restoration of ponderosa pine forests requires an understanding of the heterogeneity of contemporary and historical stand structures. We assessed spatial and temporal patterns of tree establishment, mortality and size structure over a 30-year period in an old-growth ponderosa pine stand in the mid-montane zone of the Colorado Front Range. We analyzed spatial patterns and spatial associations using Ripley's K(t) and K₁₂(t) and then modeled the patterns using point process models. Forest age structure was estimated by aging a sub-sample of trees in the stand. Climate appeared to play a significant role in the coarse-scale temporal pattern of regeneration events. Stand structure (distribution of patches, light availability, and seed trees) influenced the spatial and temporal pattern of more recent regeneration events. Patchy regeneration resulted in spatial independence and some segregation of size classes. Older trees in the stand (40–55 cm dbh) exhibited some regularity in their spatial distribution at short distances indicating that patterns of mortality had been historically patchy. Contemporary patterns of mortality were mostly patchy, and mountain pine beetles caused a significant amount of mortality in the 1970s and 1980s. Both establishment and mortality retained spatial patterns that were somewhat consistent with pre-settlement forests, despite changes in driving processes.     

Does the herb Acanthus pubescens and/or elephants suppress tree regeneration in disturbed Afrotropical forest?

As a result of extensive deforestation, the survival of many tropical forest species may depend on disturbed forests. However, recent studies demonstrate that tree regeneration following logging can be slow, decreasing the conservation value of disturbed forests. Here we test whether the dominating herb, Acanthus pubescens, in the understory of logged forest, suppresses tree regeneration in Kibale National Park, Uganda. We compare the establishment, growth, and survival of tree seedlings over 2.5 years between sites cleared of A. pubescens and controls. We also consider the role of elephants (Loxodonta africana) in inhibiting tree regeneration. Seedling abundance and species richness was greater in cleared than control plots at the start of the study. Seedling abundance decreased over the study, but remained greater in cleared than control plots. Species richness did not vary over the study. However, species composition of the cleared plots was marginally different from control plots at the start, but converged on the composition of control plots by the end of the study. This suggests a common regeneration assemblage derived from reorganisation of species, rather than recruitment of new species. There was no difference in recruitment rates between cleared and control plots. These findings suggest no strong direct effects of the herb on tree regeneration in this disturbed forest. However, when we modelled (i.e., removed) the effect of elephants on regeneration, the abundance of seedlings increased in both plot types and remained reasonably constant over time. The decline in density of regenerating stems over the study was most likely caused by elephants visiting the plots. We conclude that elephants have a far greater effect in suppressing tree regeneration at Kibale than the herb stratum and are ultimately responsible for arresting succession in disturbed forest. However, as intensive logging creates conditions favourable for A. pubescens, which is eaten by elephants, the best management intervention is to constrain logging intensities to below levels above which widespread establishment of A. pubescens is promoted, thus limiting elephant activity in regenerating areas.     

Regeneration of Rocky Mountain bristlecone pine (Pinus aristata) and limber pine (Pinus flexilis) three decades after stand-replacing fires

Rocky Mountain bristlecone pine (Pinus aristata) and limber pine (Pinus flexilis) are important high-elevation pines of the southern Rockies that are forecast to decline due to the recent spread of white pine blister rust (Cronartium ribicola) into this region. Proactive management strategies to promote the evolution of rust resistance and maintain ecosystem function require an improved understanding of the role of disturbance on the population dynamics of both species and environmental conditions that favor seedling establishment. We examined patterns of bristlecone and limber pine regeneration across the perimeters of three, 29-year-old, high-severity burns in northern, central, and southern Colorado: Ouzel, Badger Mountain, and Maes Creek, respectively. Both species exhibited a very protracted regeneration response to these fires. Bristlecone pine regeneration was concentrated near burn edges and beneath surviving seed sources. This spatial pattern is consistent with limitations incurred by wind-dispersal, also borne out by the low occurrence of seedling clusters. Relative to unburned stands, the absolute abundance of bristlecone pine generally increased only on plots retaining some surviving trees. Limber pine regeneration pattern varied between sites: high in the burn interior at Ouzel, concentrated at burn edges at Badger, and mostly in unburned stands at Maes. Clark’s Nutcracker dispersal of limber pine in each study area was indicated by high seedling distance from possible seed sources and high frequencies of clustered stems. Except at Ouzel, the absolute abundance of limber pine decreased in burns. Across sites, establishment by both species was boosted by nearby nurse objects (rocks, fallen logs, and standing tree trunks), a relationship that extended out at least as far as the closest three such objects, usually found within 50 cm. Fire decreased the frequency of Pedicularis but increased Castilleja and Ribes species (alternate hosts of white pine blister rust), though only one species, R. cereum, was positively associated with either pine species. We conclude that regeneration of bristlecone and limber pine may benefit from natural disturbance or proactive management creating appropriately sized openings and microtopographic structure (e.g., abundant fallen logs); however, beneficial responses may require many decades to be achieved.     

Role reversal in the stand dynamics of an angiosperm–conifer forest: Colonising angiosperms precede a shade-tolerant conifer in Afrotemperate forest

In mixed angiosperm–conifer forests worldwide, infrequent landscape-level catastrophic disturbances create a mosaic of persistent and different aged forest stands in the landscape with varying levels of dominance by the conifer component. In the ‘temporal stand replacement model’ (TSRM), disturbance creates conditions favouring a colonising cohort that is replaced by a suite of relatively shade-tolerant canopy species, which establish following the synchronous senescence of the pioneer canopy. In most southern hemisphere mixed angiosperm–conifer forests, with the exception of those in southern Africa, the establishing cohort is usually a large and very long-lived (550–650 years) conifer that is gradually replaced by angiosperms. As an explanation of the apparent dominance of the conifer Podocarpus latifolius, we examine the efficacy of the TSRM in mixed Afrotemperate forests where the establishing cohort is not a conifer. Forest succession in Afrotemperate forests was deterministic with the successive replacement of species determined first by their establishment success in shaded environments, and second, by their relative longevity. Several angiosperm species that were common canopy dominants established a pioneer cohort but were gradually replaced by P. latifolius, a shade-tolerant species. Continuous regeneration beneath the angiosperm canopy by P. latifolius eliminates synchronous canopy senescence, a key feature of the TSRM, as a mechanism driving the temporal replacement of canopy species. Senescing angiosperms created canopy gaps that were colonised by grasses and ferns, which suppressed canopy tree regeneration. In contrast, with continuous regeneration beneath the shaded canopy, P. latifolius gains a critical advantage over angiosperms at gap formation. Thus, in the absence of fairly large-scale natural disturbances, conifers come to dominate Afrotemperate forests. Commensurate with the latter, conifers in Podocarpus-forest were dated to approximately 320 years, more than 100 years older than the oldest P. latifolius in angiosperm-dominated forest. Tree life-history differences (shade tolerance, longevity) and the time since disturbance drive successional change from an angiosperm-dominated system to a stage dominated by P. latifolius. In general, the TSRM is a plausible explanation for the observed canopy tree structure and dynamics in mixed Afrotemperate forests. South African Afrotemperate forest is unusual among other southern hemisphere mixed angiosperm–conifer forests in that a suite of angiosperm canopy species, rather than a single conifer species, forms the colonising cohort.     

The influences of conifer succession, physiographic conditions and herbivory on quaking aspen regeneration after fire

Fire suppression over the last century has increased conifer expansion and dominance in aspen-conifer forests, which appears to be a driving force behind aspen decline in some areas. The primary objective of this study was to examine how increasing conifer dominance affects aspen regeneration vigor following the return of fire. The influence of physiographic features and herbivory on aspen regeneration vigor were also examined. The study was conducted in the Sanford fire complex located in the Dixie National Forest in southern Utah, USA, where more than 31,000 hectares burned in the summer of 2002. Seven years after the burn (at 66 locations) we measured aspen regeneration density and height as response variables and former stand composition and density (the burned trees were still standing), soil characteristics, slope, aspect and presence or absence of herbivory as independent variables. Aspen regeneration (root suckering) densities ranged from <500 to 228,000 stems/hectare with an average of 37,000 stems/hectare. Post-fire aspen regeneration density was most strongly correlated with pre-fire stand successional status (as measured by stand composition and species abundance), with percent conifer abundance (R² = −0.55) and overstory aspen density (R² = + 0.50) being the most important. Average aspen suckering densities ranged from approximately 60,000 stems/hectare in what were relatively pure aspen stands (>90% aspen) to less than 5000 stems/hectare in stands where conifer abundance was greater than 90%. Soil C, N, and P showed positive correlations (R² = 0.07 to 0.17) with aspen regeneration vigor, while soil texture had a relatively weak influence on sucker regeneration. Aspen regeneration densities were 15% lower on north facing aspects compared to east, west and south facing aspects with slope steepness showing no correlation with regeneration vigor. Regeneration density was significantly lower (8%) at sites with evidence of herbivory versus sites where herbivory was absent. Overall, the aspen regeneration response in the Sanford fire complex was strong despite high wildlife densities, which may be related to disturbance size. Where the maintenance of aspen is desired in the landscape we recommend promoting fire when the percentage of overstory conifer stems is greater than 80% or overstory aspen density is less than 200 overstory stems/hectare.     

Tree structure and diversity of lowland Atlantic forest fragments: comparison of disturbed and undisturbed remnants

This study describes the tree community structure of three moist lowland Atlantic Forest fragments in Rio de Janeiro State, southeastern Brazil. Two fragments were disturbed and an undisturbed one was used as reference. Our hypothesis was that disturbed fragments show distinct structural patterns in comparison with undisturbed stands due to past disturbance practices and forest fragmentation. Four 100 ×5 m sampling plots were demarcated in each fragment and all live and dead trees with DBH C 5 cm were located, measured and identified. The results supported our hypothesis, due to the high values found for standing dead trees, an increase of dominance of a few pioneer species, lower values of large trees and species richness in disturbed fragments in comparison with the undisturbed one. The advanced fragmentation process in the Southern Brazilian lowland areas and the high species richness in undisturbed areas highlight these forest fragments as priority areas for conservation and management.     

Relationship between annual rainfall and tree mortality in a tropical dry forest: Results of a 19-year study at Mudumalai,southern India

Variability in rainfall is known to be a major influence on the dynamics of tropical forests, especially rates and patterns of tree mortality. In tropical dry forests a number of contributing factors to tree mortality, including dry season fire and herbivory by large herbivorous mammals, could be related to rainfall patterns, while loss of water potential in trees during the dry season or a wet season drought could also result in enhanced rates of death. While tree mortality as influenced by severe drought has been examined in tropical wet forests there is insufficient understanding of this process in tropical dry forests. We examined these causal factors in relation to inter-annual differences in rainfall in causing tree mortality within a 50-ha Forest Dynamics Plot located in the tropical dry deciduous forests of Mudumalai, southern India, that has been monitored annually since 1988. Over a 19-year period (1988–2007) mean annual mortality rate of all stems >1 cm dbh was 6.9 ± 4.6% (range = 1.5–17.5%); mortality rates broadly declined from the smaller to the larger size classes with the rates in stems >30 cm dbh being among the lowest recorded in tropical forest globally. Fire was the main agent of mortality in stems 1–5 cm dbh, elephant-herbivory in stems 5–10 cm dbh, and other natural causes in stems >10 cm dbh. Elephant-related mortality did not show any relationship to rainfall. On the other hand, fire-related mortality was significantly negatively correlated to quantity of rainfall during the preceding year. Mortality due to other causes in the larger stem sizes was significantly negatively correlated to rainfall with a 2–3-year lag, suggesting that water deficit from mild or prolonged drought enhanced the risk of death but only with a time lag that was greater than similar lags in tree mortality observed in other forest types. In this respect, tropical dry forests growing in regions of high rainfall variability may have evolved greater resistance to rainfall deficit as compared to tropical moist or temperate forests but are still vulnerable to drought-related mortality.