Spatial patterns and interspecific associations of three canopy species at different life stages in a subtropical forest, China

Spatial patterns of species at different life stages are an important aspect for understanding causal mechanisms that facilitate species co-existence.Using Ripley's univariate L(t) and bivariate L12(t) functions,we analyzed the spatial patterns and interspecific associations of three canopy species at different life history stages in a 20-ha subtropical forest plot in Dinghushan Nature Reserve.Based on diameter at breast height (DBH),four life stages were distinguished.Castanopsis chinensis and Schima superba showed a unimodal DBH distribution.Engelhardtia roxburghiana showed a bimodal curve.L(t) function analysis showed significantly aggregated distributions of all three species at later life stages and random distribution at early life stages at some scales.From the analysis of L12(t) function,the results showed the positive association was a dominant pattern for most species pairs at most scales but the intensity of association decreases with the increase of life stages.Juveniles of the three species had no negative intra- and interspecific associations with the older life stages.Only premature trees were suppressed by overmature trees at some scales.Considering these results,we found three canopy-dominant species that lacked regeneration.There was no direct competition occurring between understorey individuals.Young trees can grow well under conspecific species with two other species.Longevity and lack of regeneration led to a large number of trees stored in mature and overmature stages,therefore,intra-and inter-competition can be strong at later life stages.     

不同干扰强度下针阔混交林树种空间格局及种间关联性

为了探究不同干扰程度下针阔混交林树种空间格局变化及树种种间关联性,本研究以关帝山庞泉沟自然保护区不同干扰强度的针阔混交林为对象,选取郁闭度、林分密度、伐桩数量作为划分干扰程度的因子,采用相对影响法,将林分划分为未干扰、中度干扰和严重干扰3个等级.利用空间点格局K2函数分析了林分的空间分布格局及其不同树种的种间关联.结果...     

Fire disturbance and forest structure in old‐growth mixed conifer forests in the northern Sierra Nevada,California

Question: This study evaluates how fire regimes influence stand structure and dynamics in old‐growth mixed conifer forests across a range of environmental settings. Location: A 2000‐ha area of mixed conifer forest on the west shore of Lake Tahoe in the northern Sierra Nevada, California. Methods: We quantified the age, size, and spatial structure of trees in 12 mixed conifer stands distributed across major topographic gradients. Fire history was reconstructed in each stand using fire scar dendrochronology. The influence of fire on stand structure was assessed by comparing the fire history with the age, size, and spatial structure of trees in a stand. Results: There was significant variation in species composition among stands, but not in the size, age and spatial patterning of trees. Stands had multiple size and age classes with clusters of similar aged trees occurring at scales of 113 ‐ 254 m². The frequency and severity of fires was also similar, and stands burned with low to moderate severity in the dormant season on average every 9–17 years. Most fires were not synchronized among stands except in very dry years. No fires have burned since ca. 1880. Conclusions: Fire and forest structure interact to perpetuate similar stand characteristics across a range of environmental settings. Fire occurrence is controlled primarily by spatial variation in fuel mosaics (e.g. patterns of abundance, fuel moisture, forest structure), but regional drought synchronizes fire in some years. Fire exclusion over the last 120 years has caused compositional and structural shifts in these mixed conifer forests.     

Large old trees increase growth under shifting climatic constraints: Aligning tree longevity and individual growth dynamics in primary mountain spruce forests

In a world of accelerating changes in environmental conditions driving tree growth, tradeoffs between tree growth rate and longevity could curtail the abundance of large old trees (LOTs), with potentially dire consequences for biodiversity and carbon storage. However, the influence of tree-level tradeoffs on forest structure at landscape scales will also depend on disturbances, which shape tree size and age distribution, and on whether LOTs can benefit from improved growing conditions due to climate warming. We analyzed temporal and spatial variation in radial growth patterns from ~5000 Norway spruce (Picea abies [L.] H. Karst) live and dead trees from the Western Carpathian primary spruce forest stands. We applied mixed-linear modeling to quantify the importance of LOT growth histories and stand dynamics (i.e., competition and disturbance factors) on lifespan. Finally, we assessed regional synchronization in radial growth variability over the 20th century, and modeled the effects of stand dynamics and climate on LOTs recent growth trends. Tree age varied considerably among forest stands, implying an important role of disturbance as an age constraint. Slow juvenile growth and longer period of suppressed growth prolonged tree lifespan, while increasing disturbance severity and shorter time since last disturbance decreased it. The highest age was not achieved only by trees with continuous slow growth, but those with slow juvenile growth followed by subsequent growth releases. Growth trend analysis demonstrated an increase in absolute growth rates in response to climate warming, with late summer temperatures driving the recent growth trend. Contrary to our expectation that LOTs would eventually exhibit declining growth rates, the oldest LOTs (>400 years) continuously increase growth throughout their lives, indicating a high phenotypic plasticity of LOTs for increasing biomass, and a strong carbon sink role of primary spruce forests under rising temperatures, intensifying droughts, and increasing bark beetle outbreaks.     

亚高山云冷杉混交林树木生长释放与干扰分析

对云南碧塔海亚高山云冷杉林内４个样地冠层树木的生长压制和释放的历史,用树木年轮分析方法进行了重建,然后根据生长释放频率推测林冠干扰强度（每１０年冠层树木的死亡百分率）。４个林分（１个中龄林,３个成过熟林）生长释放的平均百分率为４８％～９２％。中龄林内,平均生长释放频率为７１％／１０ａ,成过熟林则为７４％～９５％／１０ａ,在油麦吊云杉〔Ｐｉｃｅａｂｒａｃｈｙｔｙｌａｖａｒ．ｃｏｍｐｌａｎａｔａ（Ｍａｓｔ．）ＣｈｅｎｇｅｘＲｅｈｄ．〕占优势的林分和大果红杉（Ｌａｒｉｘｐｏｔａｎｉｎｉｖａｒ．ｍａｃｒｏｃａｒｐａＬａｗ）油麦吊云杉混交林分内,估测的林冠干扰强度分别为４８％／１０ａ和５９％／１０ａ。     

Fire disturbance and forest structure in an old‐growth Pinus ponderosa forest,southern Cascades,USA

Questions: Did fire regimes in old‐growth Pinus ponderosa forest change with Euro‐American settlement compared to the pre‐settlement period? Do tree age structures exhibit a pattern of continuous regeneration or is regeneration episodic and related to fire disturbance or fire‐free periods? Are the forests compositionally stable? Do trees have a clumped spatial pattern and are clumps even‐ or mixed‐age? How might information from this old‐growth forest inform current restoration and management practices? Location: A 235‐ha old‐growth forest in the Ishi Wilderness, southern Cascade Mountains, California. Methods: Age, size, and spatial pattern of trees were quantified in seven stands. Fire history was reconstructed using fire scar dendrochronology. The influence of fire on stand structure was assessed by comparing fire history with age, size, and spatial structure of trees and identifying and measuring trees killed by two recent fires. Results: Species composition in plots was similar but density and basal area of tree populations varied. Age structure for P. ponderosa and Quercus kelloggii showed periods of episodic recruitment that varied among plots. Fire disturbance was frequent before 1905, with a median period between fires of 12 years. Fire frequency declined after 1905 but two recent fires (1990, 1994) killed 36% and 41% of mostly smaller diameter P. ponderosa and Q. kelloggii. Clusters of similar age trees occurred at scales of 28‐1018 m² but patches were not even‐aged. Interactions between tree regeneration and fire promoted development of uneven age groups of trees. Conclusions: Fire disturbance strongly influenced density, basal area, and spatial structure of tree populations. Fire exclusion over the last 100 years has caused compositional and structural changes. Two recent fires, however, thinned stands and created gaps favorable for Q. kelloggii and P. ponderosa regeneration. The effects of infrequent 20th century fire indicate that a low fire frequency can restore and sustain structural characteristics resembling those of the pre‐fire suppression period forest.     

Effects of fire and spruce beetle outbreak legacies on the disturbance regime of a subalpine forest in Colorado

Aim There is increasing research attention being given to the role of interactions among natural disturbances in ecosystem processes. We studied the interactions between fire and spruce beetle (Dendroctonus rufipennis Kirkby) disturbances in a Colorado subalpine forest. The central questions of this research were: (1) How does fire history influence stand susceptibility to beetle outbreak? And conversely, (2) How does prior occurrence of a beetle outbreak influence stand susceptibility to subsequent fire? Methods We reconstructed the spatial disturbance history in a c. 4600 ha area by first identifying distinct patches in the landscape on aerial photographs. Then, in the field we determined the disturbance history of each patch by dating stand origin, fire scars, dates of mortality of dead trees, and releases on remnant trees. A geographical information system (GIS) was used to overlay disturbance by fire and spruce beetle. Results and main conclusions The majority of stands in the study area arose following large, infrequent, severe fires occurring in c. 1700, 1796 and 1880. The study area was also affected by a severe spruce beetle outbreak in the 1940s and a subsequent low‐severity fire. Stands that originated following stand‐replacing fire in the late nineteenth century were less affected by the beetle outbreak than older stands. Following the beetle outbreak, stands less affected by the outbreak were more affected by low‐severity fire than stands more severely affected by the outbreak. The reduced susceptibility to low‐severity fire possibly resulted from increased moisture on the forest floor following beetle outbreak. The landscape mosaic of this subalpine forest was strongly influenced by the interactions between fire and insect disturbances.     

Microsite variation and climate result in stand dynamics variation in pristine Pinus sylvestris mires: Evidence from a stand structure study

Questions: How do climate conditions and the site's ecohy‐ drological properties affect the age and size structure of natural Pinus sylvestris stands on pristine boreal mires? How do the long‐term stand dynamics on mires proceed as stands age? Do the mire stands reach a balanced, old‐growth stage? Location: Boreal mire forests in southern and northern Finland. Methods: Tree age and diameter distributions were analysed in 52 stands in two climate areas and in two mire site types with different ecohydrological properties. Temporal stand dynamics were examined by (1) comparing the graphs of the stands’ mean tree ages by diameter at breast height (1.3 m) classes and (2) describing the changes in stand characteristics and stand age and size structures as a function of stand dominant age in a chronosequence. Results: In the south, the DBH distributions were mostly unimodal and bell‐shaped in both site type groups. Age distributions were multimodal and flat in fully‐stocked sites but more uneven in sparsely forested composite sites. In the north, both the age and size distributions were clearly uneven in both site type groups. Tree age and size variation increased with stand age, but levelled out in the long term. Particularly in the south, the abundance of small trees decreased as stand age increased. Conclusions: The pine stands on pristine boreal mires are more dynamic than anticipated and are generally not characterised by a balanced, self‐perpetuating structure. Their dynamics reflect differences in climate and ecohydrology: on stocked sites in favourable boreal conditions, the stands showed structures typically resultant of inter‐tree competition processes that control tree growth and regeneration, whereas in harsh boreal climates, the tree regeneration process is ongoing diversifying the stand structure.     

Effects of disturbance and size structure on the regeneration process in a sub-boreal coniferous forest,northern Japan

The stand structure and disturbance history in a sub-boreal coniferous forest dominated byPicea jezoensis, Picea glehnii andAbies sachalinensis were investigated in four study plots set up in Taisetsuzan National Park, Japan. The effect of stand characteristics on the growth and mortality rates of understory trees was examined. Although all the stands showed inverse J-shape d.b.h. (diameter at breast height) distributions, the age structure and disturbance history differed amongst the stands. The stands with wide d.b.h. distribution (i.e. large CV and skewness) were more uneven-aged than those with narrow d.b.h. distribution (i.e. small CV and skewness). The disturbance-return interval based on the model of Hett and Loucks was 31 to 65 years. The gap ratio in the canopy was also different among the stands. These suggest that the variations in stand structure represent different occurrences of natural disturbances. Furthermore, the structural features such as size structure, canopy gap ratio and density of canopy trees also affected the growth dynamics of understory trees (≥2 m in height and <10 cm in diameter at breast height). The growth and mortality rates of understory trees changed with the canopy gap ratio and canopy tree density. The understory trees of stands with wide canopy d.b.h. distribution had higher growth and canopy recruitment rates than those of stands with narrow canopy d.b.h. distribution, contributing to the maintenance of continuous stand stratification. The understory trees of stands with narrow canopy d.b.h. distribution showed lower growth and higher mortality rates than those of stands with narrow canopy d.b.h. distribution, leading to the formation of a single-canopy structure. It is suggested that natural disturbance governs the regeneration process in the future by affecting the growth and mortality patterns of understory trees through the stand structure (size and age structure, canopy tree density, canopy gap ratio).     

Structure and stand development in three subalpine Norway spruce (Picea abies (L.) Karst.) stands in Paneveggio (Trento, Italy)

1. Three permanent plots (100×0 m) were established in the subalpine Norway spruce (Picea abies (L.) Karst.) forest of Paneveggio in the spring of 1993, to begin a long-term forest ecosystem research project. The main purpose of these plots was to provide information about subalpine Norway spruce stand dynamics and to provide suggestions for close-to-nature silviculture. 2. The three stands were selected to represent the most common forest structures in the Paneveggio forest. The first stand is close to forestry roads, has a relatively regular and continuous canopy, and thinning and cutting operations only ended in the 1980s; the second stand is far from forest roads and has developed without anthropogenic influence for several decades; the third one is located at the present upper limit of the pure spruce forest and, apparently, was heavily used in the past as a pasture. 3. The first step in the investigation was to describe the structure and to study the history of the three stands using both written evidence from manage- ment plans and biological archives from tree rings. 4. The stands in plots 1 and 2 began to establish after a disturbance that removed part of the previous stands according to dendroecological studies, which are partially supported by written evidence. The remaining parts of these stands were eliminated by two major disturbances that occurred during the following decades. Written records about the use of the forest lead us to assume that the initial disturbances that occurred in the two stands were logging activities as a part of a group shelterwood system. The stand in plot 2 has developed without significant human interference for about half a century as confirmed by the presence of many dead trees. The stand in plot 3 consists of old trees that were part of an open stand and a secondary population that established after cessation of grazing. 5. The study has confirmed that dendroecological techniques can be used to identify occurrence and intensity of previous disturbance in forests stands, although at Paneveggio it is difficult to distinguish between natural and anthropogenic disturbances in the tree ring record. The presence of human activity necessitates investigation of multiple lines of evidence. 6. Paneveggio's forest management plans were useful in the interpretation of the data obtained through dendroecological analysis, although events did not always correspond because data from the management plans (yearly thinning, felling, wind-throw damage) never gave stand-level details, but applied to areas of several hectares. Despite these limitations, the information included in the management plans is of crucial importance in studying stand history and only by using all these sources of information is it possible to delineate the most important features of the history and disturbance that affected the origin and subsequent growth of the forest stands.     

Dominant species and dispersal limitation regulate tree species distributions in a 20‐ha plot in Xishuangbanna,southwest China

Habitat heterogeneity and dispersal limitation are widely considered to be the two major mechanisms in determining tree species distributions. However, few studies have quantified the relative importance of these two mechanisms at different life stages of trees. Moreover, rigorous quantification of the effects of dominant tree species in determining species distributions has seldom been explored. In the present study, we tested the hypothesis that the distribution of tree species is regulated by different mechanisms at different life history stages. In particular, we hypothesised that dispersal limitation regulates the distribution of trees at early life stages and that environmental factors control the distribution of trees as they grow, because of niche differentiation resulting from environmental filtering. To test this, trees in 400‐m² quadrats in a 20‐ha plot in Xishuangbanna, southwest China were grouped into four classes on the basis of the diameter at breast height (DBH) that roughly represent different stages in the life history of trees. A neighbourhood index was computed to represent a neutral spatial autocorrelation effect. We used both biotic (dominant species) and abiotic (topography and soil) predictor variables to model the distribution of each target species while controlling for spatial autocorrelation within each of the DBH classes. To determine which factors played the largest role in regulating target species distribution, the simulated annealing method was used in model selection based on Akaike information criterion (AIC) values. The results showed that the relative importance of neutral and niche processes in regulating species distribution varied across life stages. The neutral neighbourhood index played the most important role in determining the distributions of small trees (1 cm ≤ DBH ≤ 10 cm), and dominant species, as biotic environmental predictor variables, were the next most important regulators for trees of this size. Environmental predictor variables played the most important role in determining the distributions of large trees (10 cm ≤ DBH). This finding builds on previous research into the relative importance of neutral and niche processes in determining species distributions regardless of life stages or DBH classes.     

Large‐scale disturbance legacies and the climate sensitivity of primary Picea abies forests

Determining the drivers of shifting forest disturbance rates remains a pressing global change issue. Large‐scale forest dynamics are commonly assumed to be climate driven, but appropriately scaled disturbance histories are rarely available to assess how disturbance legacies alter subsequent disturbance rates and the climate sensitivity of disturbance. We compiled multiple tree ring‐based disturbance histories from primary Picea abies forest fragments distributed throughout five European landscapes spanning the Bohemian Forest and the Carpathian Mountains. The regional chronology includes 11,595 tree cores, with ring dates spanning the years 1750–2000, collected from 560 inventory plots in 37 stands distributed across a 1,000 km geographic gradient, amounting to the largest disturbance chronology yet constructed in Europe. Decadal disturbance rates varied significantly through time and declined after 1920, resulting in widespread increases in canopy tree age. Approximately 75% of current canopy area recruited prior to 1900. Long‐term disturbance patterns were compared to an historical drought reconstruction, and further linked to spatial variation in stand structure and contemporary disturbance patterns derived from LANDSAT imagery. Historically, decadal Palmer drought severity index minima corresponded to higher rates of canopy removal. The severity of contemporary disturbances increased with each stand's estimated time since last major disturbance, increased with mean diameter, and declined with increasing within‐stand structural variability. Reconstructed spatial patterns suggest that high small‐scale structural variability has historically acted to reduce large‐scale susceptibility and climate sensitivity of disturbance. Reduced disturbance rates since 1920, a potential legacy of high 19th century disturbance rates, have contributed to a recent region‐wide increase in disturbance susceptibility. Increasingly common high‐severity disturbances throughout primary Picea forests of Central Europe should be reinterpreted in light of both legacy effects (resulting in increased susceptibility) and climate change (resulting in increased exposure to extreme events).     

Forest development in canopy gaps in old-growth beech (Nothofagus) forests,New Zealand

Abstract. Tree size and age structures, treefall and canopy gap characteristics, and regeneration responses to treefalls were compared for three stands of old-growth beech (Nothofagus) forest dominated by N. fusca and N. menziesii on the South Island, New Zealand. Treefall gaps (up to 1000 m²) were most often caused by standing trees killed by drought and/or insect attack, or by trees snapped by wind. The causes of gap formation and the size and age distributions of treefall gaps varied between localities because of spatial and temporal differences in the histories of disturbance. At Fergies Bush where drought-related dieback had produced many large gaps with standing dead trees, gaps were generally young. Conversely, at Station Creek, small, old gaps formed by bolesnap dominated the disturbance regime. At Rough Creek, gaps of all ages and sizes were found along with an almost complete fern cover, and abundant shrubs and occasional subcanopy hardwood trees. Although overall patterns of regeneration were unrelated to differences in gap size, the relative abundance of N. fusca and N. menziesii varied between localities according to the seemingly minor differences in forest structure and disturbance history described above. Interpretations of regeneration response to gap parameters, therefore, need to account for differences in disturbance history between sites. Differences in the disturbance history between localities will also influence rates of gap closure, and because closure rates are used to estimate forest turnover times, meaningful comparisons of disturbance regimes for different forest types can only be made if this intersite variability is addressed.     

Stand Structure and Maintenance of <Emphasis Type="Italic">Picea jezoensis</Emphasis> in a Northern Temperate Forest,South Korea

Stand structure and spatial distribution of Picea jezoensis (Siebold et Zucc.) Carrière on Mt. Gyebang, Korea was investigated to provide information on the structural characteristics and the maintenance of P. jezoensis population in northern temperate mixed coniferous forests. Height and diameter at breast height (DBH) distribution, age, growth, and spatial distribution patterns of P. jezoensis were examined in thirty nine 100-400 m² quadrats or circular plots. The overall stand structure attributes in the study sites are stem density of 709 trees ha⁻¹, a mean DBH of 12.8 cm, and a mean height of 5.6 m, with reverse J shapes of DBH and height distributions. The stem density of P. jezoensis population was 81 trees ha⁻¹, a mean DBH of 20.7 cm, and a mean height of 9.1 m, showing bimodal-like shapes in age and DBH distributions. Several growth release periods implied that P. jezoensis stands experienced small disturbances. The radius of patches of similar-sized P. jezoensis in the variogram was equivalent with the height of the tallest trees, indicating that patches were established following the fall of trees in the upper canopy layer. Small windthrows in this region contributed to the maintenance of the P. jezoensis stand by releasing sapling growth and providing nursing logs and space for seedlings.     

Changing growth response to wildfire in old‐growth ponderosa pine trees in montane forests of north central Idaho

North American fire‐adapted forests are experiencing changes in fire frequency and climate. These novel conditions may alter postwildfire responses of fire‐adapted trees that survive fires, a topic that has received little attention. Historical, frequent, low‐intensity wildfire in many fire‐adapted forests is generally thought to have a positive effect on the growth and vigor of trees that survive fires. Whether such positive effects can persist under current and future climate conditions is not known. Here, we evaluate long‐term responses to recurrent 20th‐century fires in ponderosa pine, a fire‐adapted tree species, in unlogged forests in north central Idaho. We also examine short‐term responses to individual 20th‐century fires and evaluate whether these responses have changed over time and whether potential variability relates to climate variables and time since last fire. Growth responses were assessed by comparing tree‐ring measurements from trees in stands burned repeatedly during the 20th century at roughly the historical fire frequency with trees in paired control stands that had not burned for at least 70 years. Contrary to expectations, only one site showed significant increases in long‐term growth responses in burned stands compared with control stands. Short‐term responses showed a trend of increasing negative effects of wildfire (reduced diameter growth in the burned stand compared with the control stand) in recent years that had drier winters and springs. There was no effect of time since the previous fire on growth responses to fire. The possible relationships of novel climate conditions with negative tree growth responses in trees that survive fire are discussed. A trend of negative growth responses to wildfire in old‐growth forests could have important ramifications for forest productivity and carbon balance under future climate scenarios.     

Hurricane Disturbance Alters Secondary Forest Recovery in Puerto Rico

Land-use history and large-scale disturbances interact to shape secondary forest structure and composition. How introduced species respond to disturbances such as hurricanes in post-agriculture forest recovery is of particular interest. To examine the effects of hurricane disturbance and previous land use on forest dynamics and composition, we revisited 37 secondary forest stands in former cattle pastures across Puerto Rico representing a range of exposure to the winds of Hurricane Georges in 1998. Stands ranged from 21 to>80 yr since agricultural abandonment and were measured 9 yr posthurricane. Stem density decreased as stands aged, while basal area and species richness tended to increase. Hurricane disturbance exerted contrasting effects on stand structure, contingent on stand age. In older stands, the basal area of large trees fell, shifting to a stand structure characteristic of younger stands, while the basal area of large trees tended to rise in younger stands with increasing hurricane disturbance. These results demonstrate that large-scale natural disturbances can alter the successional trajectory of secondary forest stands recovering from human land use, but stand age, precipitation and soil series were better predictors of changes in stand structure across all study sites. Species composition changed substantially between census intervals, but neither age nor hurricane disturbance consistently predicted species composition change. However, exposure to hurricane winds tended to decrease the abundance of the introduced tree Spathodea campanulata, particularly in smaller size classes. In all sites the abundance of the introduced tree Syzygium jambos showed a declining trend, again most strongly in smaller size classes, suggesting natural thinning through succession.     

Effects of environment and grazing disturbance on tree establishment in meadows of the central Cascade Range,Oregon, USA

Abstract. Within the last century there has been widespread establishment of trees in mountain meadows of the Pacific Northwest. We reconstructed patterns of tree invasion at 17 meadow sites in the central Cascade Range of Oregon, USA -sites representing diverse physical environments and vegetation types and experiencing different histories of recent anthropogenic disturbance (sheep grazing). Spatial distributions and age structures of invasive tree populations were analysed with respect to climatic records and grazing history. Patterns of establishment varied considerably among meadows, reflecting strong differences in environment and grazing history. In montane hydric meadows, tree establishment was spatially clumped beneath large old trees and on elevated microsites; however the timing of invasion differed between sites with stable versus fluctuating water tables. In upland mesic/dry montane meadows, timing of invasion corresponded with cessation of sheep grazing (early 1940s) and the onset of wetter summers (mid 1940s). In the subalpine zone, climate and aspect interacted to produce contrasting histories of invasion on north- and south-facing slopes. Establishment on north-facing slopes, concentrated in heath-shrub communities, coincided with regional warming (ca. 1920–1945) when snowpacks were lighter and melted earlier. Recruitment of trees onto south-facing slopes occurred later, when conditions were wetter (1945–1985). In many environments, the spatial distribution of recruitment suggests that once trees have established, autogenic factors become increasingly important as individual trees or groups of trees alter the physical or biotic conditions that once inhibited establishment. Knowledge of the factors that influence invasion, and of their varying importance across gradients in environment and vegetation, is critical to predicting future changes in these dynamic systems.     

Contrasting patterns of tree mortality in late‐successional Picea abies stands in two areas in northern Fennoscandia

Question: What are tree mortality rates and how and why do they vary in late‐successional Picea abies‐dominated forests? Do observed tree mortality patterns allow comparative assessment of models of long‐term stand development? Location: Northern boreal Fennoscandia. Methods: We measured stand structure in 10 stands in two different areas. We determined age distributions and constructed a chronology of tree deaths by cross‐dating the years of death of randomly sampled dead trees. Results: The stands in the two areas had contrasting tree age distributions, despite similar live tree structure. In one area, stands were relatively even‐aged and originated following a stand‐replacing fire 317 years earlier. The stands in the second area had an uneven age structure and virtually no signs of past fires, suggesting a very long period since the last major disturbance. The younger stands were characterized by a high mortality rate and inter‐annual variation, which we attributed to senescence of the relatively even‐aged stands approaching the maximum age of P. abies. In contrast, the tree mortality rates in the older stands were low and relatively stable. Conclusions: Patterns of tree mortality were, to a large extent, dependent on the time since the last stand‐replacing disturbance, suggesting that northern boreal P. abies stands eventually reach a shifting mosaic state maintained through small‐scale dynamics, but the time needed to reach this state appears to be lengthy; even 300 years after a forest fire stands showed changes in patterns of tree mortality that were related to the developmental stage of the stands.     

Linking radial growth patterns and moderate‐severity disturbance dynamics in boreal old‐growth forests driven by recurrent insect outbreaks: A tale of opportunities,successes, and failures

In boreal landscapes, emphasis is currently placed on close‐to‐nature management strategies, which aim to maintain the biodiversity and ecosystem services related to old‐growth forests. The success of these strategies, however, depends on an accurate understanding of the dynamics within these forests. While moderate‐severity disturbances have recently been recognized as important drivers of boreal forests, little is known about their effects on stand structure and growth. This study therefore aimed to reconstruct the disturbance and postdisturbance dynamics in boreal old‐growth forests that are driven by recurrent moderate‐severity disturbances. We studied eight primary old‐growth forests in Québec, Canada, that have recorded recurrent and moderately severe spruce budworm (Choristoneura fumiferana [Clem.]) outbreaks over the 20th century. We applied an innovative dendrochronological approach based on the combined study of growth patterns and releases to reconstruct stand disturbance and postdisturbance dynamics. We identified nine growth patterns; they represented trees differing in age, size, and canopy layer. These patterns highlighted the ability of suppressed trees to rapidly fill gaps created by moderate‐severity disturbances through a single and significant increase in radial growth and height. Trees that are unable to attain the canopy following the disturbance tend to remain in the lower canopy layers, even if subsequent disturbances create new gaps. This combination of a low stand height typical of boreal forests, periodic disturbances, and rapid canopy closure often resulted in stands constituted mainly of dominant and codominant trees, similar to even‐aged forests. Overall, this study underscored the resistance of boreal old‐growth forests owing to their capacity to withstand repeated moderate‐severity disturbances. Moreover, the combined study of growth patterns and growth release demonstrated the efficacy of such an approach for improving the understanding of the fine‐scale dynamics of natural forests. The results of this research will thus help develop silvicultural practices that approximate the moderate‐severity disturbance dynamics observed in primary and old‐growth boreal forests.     

Impact of a strong typhoon on the structure and dynamics of an old-growth beech (<Emphasis Type="Italic">Fagus crenata</Emphasis>) forest,southwestern Japan

Typhoon no. 19 of 1991 (T9119) caused multiple treefalls and created large openings in an old-growth beech (Fagus crenata) forest at Mt. Daisen, in the Daisen Forest Reserve, southwestern Japan. The area of the largest opening was about 1.7 ha (300 m by 70 m). To predict the dynamics of the beech stand after the disturbance of T9119, we investigated the damage to the stand and the density and growth rate of trees with DBH=5–10 cm in a 1-ha plot covering a large part of the largest opening and the adjacent closed canopy. The beech did not regenerate immediately. The regeneration and growth rate of trees with DBH=5–10 cm were related to the frequency of the typhoon attack for at least the past century. In beech forests, small gap formation is the prevailing mode of disturbance. Our results indicate that typhoons affect the structure and dynamics of this beech stand. We suggest that both small gap formation and large-scale disturbance are important for the maintenance of beech forest in some areas.