择伐对吉林蛟河阔叶红松林群落结构及动态的影响

科学的森林经营能够优化林分结构,是调控森林生产力和生物多样性的有效手段。择伐作为森林经营的重要方式之一,其对森林结构以及群落动态的影响一直未有定论,因此迫切需要利用更加全面的数据对择伐及伐后林分特征的变化进行长期监测。根据森林大样地建立规范,2010年在吉林蛟河建立了42hm~2阔叶红松林动态监测样地,2011年冬季截取部分面积进行择伐经营,以经营样地为研究对象,运用数值变量描述采伐活动并分析择伐前后群落结构的变化;同时结合2015年的二次调查数据,以立地条件基本一致的对照样地为参照,比较林分水平和物种水平上死亡率、更新率的差异,并利用线性混合效应模型探究择伐活动对个体径向生长的影响。研究结果显示:经营样地的择伐强度为5.4%,受采伐干扰影响较大的物种主要包括色木槭、白牛槭、裂叶榆、胡桃楸、千金榆、水曲柳以及紫椴,采伐主要集中于林冠层树种,亚林层和灌木层个体很少涉及。择伐前后物种组成、径级结构等并未发生明显改变。5年间,经营样地和对照样地的林分密度都降低,对比对照样地,经营样地的死亡率较低,但其更新状况并未优于对照样地。从胸高断面积来看,经营样地整体的年平均生长量高于对照样地,表明择伐导致的稀疏对个体生长和存活起到了一定的促进作用。将采伐强度纳入线性混合效应模型中分析发现,胸径始终是影响个体生长的最重要因素,其次是树木个体之间的非对称竞争;采伐所涉及到的7个主要树种的年平均生长量均高于对照样地,但仅有紫椴的径向生长表现出对采伐干扰的显著响应。综合来看,低强度择伐对群落结构和动态的影响较小,不同物种的径向生长对择伐的响应存在一定差异。     

A comparison of structural characteristics between old-growth and postfire second-growth hemlock–hardwood forests in Adirondack Park, New York, U. S. A.

1 This study compares the structural characteristics of 12 old‐growth and six postfire second‐growth hemlock–northern hardwood stands in north central Adirondack Park, New York, in order to test the null hypothesis that there are no differences in species composition, size structure, age structure and attributes such as dead wood and canopy gaps between old‐growth stands and this type of second‐growth forest. 2 The second‐growth forests of this study regenerated following widespread logging‐related fires in either 1903 or 1908; the old growth and second growth have similar environmental settings. 3 Estimates of stand ages, derived from an increment core of the oldest tree in each stand, range from 88 to 390 years. 4 Structural attributes are related to stand age (i.e. stage of development). In comparison with the second‐growth forests of this study, older stands are characterized as (a) a larger average diameter of canopy trees; (b) a greater basal area of trees; (c) a lower density of canopy trees and of all trees ≥ 10 cm d.b.h.; (d) a higher density of eastern hemlock (Tsuga canadensis (L.) Carrière) trees; (e) a higher density of large trees (≥ 50 cm d.b.h.); (f) larger canopy gaps; and (g) a greater volume of coarse woody debris (both logs ≥ 20 cm d.b.h. and snags ≥ 10 cm d.b.h.). 5 Despite differences between old growth and second growth, especially in species composition, it appears from observations of the 18 stands that second‐growth forests are developing some structural characteristics of old growth. 6 Structural attributes of the old‐growth forests are similar to characteristics of the same forest type in geographically distant areas in eastern USA.     

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.     

Managing red pine stand structure to mitigate drought impacts

Reducing forest stand density through silvicultural thinning has demonstrated potential to mitigate drought impacts on growth; however, less has been studied on how changes in stand structure created by different thinning methods influence forest growth responses to drought. This research examined the growth responses to drought of natural-origin red pine in a long-term study contrasting thinning methods. Dendrochronological methods were used to examine growth responses during several drought events among stands where different thinning methods have been applied since 1950. Growth responses to drought were expressed as resistance (maintaining growth during drought), and resilience (regaining pre-drought growth). Results indicate that periodic thinning from above, which resulted in smaller diameters, has the potential to moderate drought-induced growth reductions. Larger tree diameters negatively influenced tree-level resistance and resilience across all treatments; however, the proportion of dominant trees in a stand had contrasting effects on stand-level drought responses. Stands thinned from above exhibited more complex vertical structure and increased stand-level resistance and resilience to drought-induced growth declines because competition is more stratified among smaller diameter trees. Opposite trends were observed in stands thinned from below, where the larger diameters and monolayered structure create greater competition among trees of similar size and crown position. The results of this study highlight the utility in managing for greater structural diversity to mitigate the negative effects of drought in red pine forest ecosystems.     

Competition and regeneration in quaking aspen–white fir (Populus tremuloides–Abies concolor) forests in the Northern Sierra Nevada,USA

Question: How does competition between quaking aspen (Populus tremuloides) and white fir (Abies concolor) affect growth and spatial pattern of each species? Location: The northern Sierra Nevada, California, USA. Methods: In paired plots in mixed aspen‐ (n=3) or white fir‐dominated (n=2) stands, we mapped trees and saplings and recorded DBH, height, species, and condition and took increment cores. We tallied seedlings by species. Tree ring widths were used as a measure of basal area change over the last decade, and canopy openness was identified using hemispherical photographs. Linear mixed models were used to relate neighborhood indices of competition, stand, and tree‐level variables to diameter increment. Spatial patterns of stems were identified using the Neighborhood Density Function. Results: White fir radial growth was higher in aspen‐ than white fir‐dominated plots. Individual‐level variables were more important for white fir than for aspen growth, while variables representing competitive neighborhood were important only for aspen. The forest canopy was more open in aspen‐ than white fir‐dominated stands, but ample aspen seedlings were observed in all stands. Canopy stems of aspen and white fir were randomly distributed, but saplings and small trees were clumped. Aspen saplings were repelled by canopy aspen stems. Conclusions: Variation in canopy openness explained more stand–stand variation in white fir than aspen growth, but high light levels were correlated with recruitment of aspen seedlings to the sapling class. Radial growth of aspen was predicted by indices of neighborhood competition but not radial growth of white fir, indicating that spacing and stem arrangement was more important for aspen than white fir growth. Fire suppression has removed a major disturbance mechanism that promoted aspen persistence and reduced competition from encroaching conifers, and current forests favor species that regenerate best by advance regeneration (white fir).     

Tree mortality and effects of release from competition in an old‐growth Fagus‐Abies‐Picea stand

Abstract. In a montane mixed Fagus‐Abies‐Picea forest in Babia Gora National Park (southern Poland), the dynamics of an old‐growth stand were studied by combining an 8‐yr annual census of trees in a 1‐ha permanent sample plot with radial increments of Abies and Picea growing in the central part of the plot. The mortality among the canopy trees was relatively high (10% in 8 yr), but the basal area increment of surviving trees slightly exceeded the losses caused by tree death. DBH increment was positively correlated with initial diameter in Abies and Picea, but not in Fagus. For individual trees smaller than the median height, basal area increment was positively related to the basal area of old snags and the basal area of recently deceased trees in their neighbourhood, but negatively related to the basal area of live trees. Dendrochronological analysis of the past growth patterns revealed numerous periods of release and suppression, which were usually not synchronized among the trees within a 0.3 ha plot. The almost normal distribution of canopy tree DBH and the small number of young individuals in the plot indicated that stand dynamics were synchronized over a relatively large area and, hence, were consistent with the developmental phase concept. On the other hand, the lack of synchronization among periods of growth acceleration in individual mature Abies and Picea trees conforms more closely to the gap‐dynamics paradigm.     

Regional tectonics, disturbance, and population size of isolated stands of Nothofagus fusca (Nothofagaceae) in a forest ecotone in south-western New Zealand

Aim New Zealand's cool temperate forests are usually dominated by one or more of the five native taxa of Nothofagus (Nothofagaceae; southern beech), but in certain regions there are sharp boundaries against podocarp–broadleaved forest where Nothofagus is rare or completely absent, either for historical (Pleistocene Glaciation) or climatic/biological (mild superhumid climate and competition) reasons. The dynamics of a Nothofagus boundary was investigated by monitoring disturbance-initiated establishment of isolated stands of N. fusca at the extreme limits of its regional distribution. Location The research was carried out in a regional forest ecotone between Nothofagus forest and podocarp–broadleaved forest in the upper Taramakau Valley, South Island, New Zealand. The survey region straddles a major, active fault system and associated tectonic movements and earthquakes with more distant epicentres have contributed to intermittent canopy disturbance of the local forests. Methods Isolated stands of Nothofagus fusca beyond the limits of continuous Nothofagus forest were investigated during two field surveys, separated by 7–10 years. Changes in population size, stem diameter of individual trees, stand basal area and mean annual diameter increment were calculated for each of fifty-four isolated stands. Types of past and recent disturbance and the probable cause of mortality of trees were noted. Results The total population of fifty-four sample stands, ranging in size from one to > 400 stems, increased by 37.4%, and compound basal area increased by 4.7% between the two surveys. Mean stem diameter growth of isolated stands was lower than expected by empirical data for N. fusca, suggesting reduced wood increment at the limits of its distribution. Tree mortality was 0.8% per year. Fifty-one per cent of the dead stems had died as a consequence of various types of natural disturbance, uproots being more common than snaps and crown breakage. Main conclusions The isolated N. fusca stands preferentially occupy sites likely to experience intermittent disturbance, mostly including disturbance of the soil cover, which facilitates their initial establishment and persistence. Because of causal relationships between mass movement on steep slopes and erosion/deposition of talus fans and river terraces, disturbance-initiated changes in forest composition are observed across a range of different landforms.     

Selection of mature and old stands by Corsican Nuthatch Sitta whiteheadi in harvested forests

In harvested forests, the bird community is largely determined by stand structure, which itself is determined by forestry practices. This study aimed to identify habitat variables determining the presence of Corsican Nuthatch Sitta whiteheadi – a threatened island endemic – in harvested Corsican Pine Pinus nigra laricio woods, with the aim of mitigating the impact of timber harvest on the bird. Comparison of occupied and unoccupied plots showed that this bird is found mostly in pure Corsican Pine stands, and is absent when more than 50% of trees are not this species. Nests were built in decaying pine snags between 20 and 100 cm diameter at breast height (dbh), but birds avoided stands with live pines < 70 cm dbh, and selected stands with pines > 80 cm dbh. Conservation of Corsican Nuthatch therefore depends on maintaining harvest rotations of more than 200 years, reducing the size of felling coupes in clear‐cutting systems or, preferably, practising selective cutting, maintaining a sufficient density of old trees and snags, and checking the encroachment of other tree species into Corsican Pine stands.     

Restoration of Old Forest Features in Coast Redwood Forests Using Early‐stage Variable‐density Thinning

To accelerate development of old forest features in coast redwood, two thinning treatments and an unthinned control were compared in three treatment areas in north coastal California. One thinning treatment was designed to restore old forest densities of 125 trees/ha and the other 250 trees/ha representing a one‐step and partial treatments to the desired stand density. Four years after treatment, numbers of trees had increased in the thinning treatments due to recruitment of new trees, but had decreased in the control due to self‐thinning. Residual trees increased in stem volume following thinning by 128% in low‐density thinning compared to 70% in the controls indicating thinning accelerated stand development. The thinning treatments also moved the species composition of these stands to a greater proportion of redwood. Considerable slash was produced by the thinning treatments but was decomposing rapidly. Black bears damaged approximately 15% of all trees and more than 38% of residual trees in the thinned treatments compared to less than 2% of all trees in the control. This damage included killing some trees and damaging other trees that survived. Decisions over restoration densities in these stands are complicated by prolonged stand development, and balancing risks and costs. In this case, the bears represent a stochastic factor that dramatically increases risk. Thinning appears to be an effective means of enhancing old forest development by accelerating tree growth, modifying species composition, and increasing stand‐level variability. Continued monitoring will be necessary to evaluate long‐term trends in density relative to effects of bear damage.     

Development processes and growth pattern of Pinus densiflora stands in central eastern Korea

Stand growth and developmental processes were investigated in Pinus densiflora Siebold et Zucc. stands of different ages in the central eastern region of Korea. Stands were inventoried and five trees per stand were sampled for stem analysis, age estimation, and growth analysis. More than 80% of sampled trees in a stand were established within 3–5 years, and most stands had a single cohort structure. The initial growth of pine seedlings was slow, but the height growth accelerated beyond 2–3 m height, 5–10 years after establishment. Linear growth was maintained until 10–12 m height, at which suppressed trees fell behind and might die out. The young stand was composed of pure pines, while few pine seedlings and saplings were found in the understory of older stands. The peak of diameter growth rate occurred around 5–15 years after tree establishment, implying that competition begins during that period. The pine stand development follows four stages: (1) the young stage when the growth rate increases and peaks; (2) the height competition stage when trees focus on height growth for light while maintaining a narrow DBH and height distribution; (3) the differentiation stage when suppressed trees die out, and the DBH distribution becomes wider; and (4) the mature stage when stands have a multi-canopy structure with a wide DBH and height distribution, while the understory is dominated by other tree species. The changes in growth rates and stand structure through forest development would be implemented to predict alterations of above-ground carbon sequestration rates.     

The response of mammals to forest fire and timber harvest in the North American boreal forest

1. This paper reviews and compares the effects of forest fire and timber harvest on mammalian abundance and diversity, throughout successional time in the boreal forest of North America. 2. Temporal trends in mammal abundance and diversity are generally similar for both harvested and burned stands, with some differences occurring in the initiation stage (0–10 years post disturbance). 3. Small mammals and ungulates are most abundant immediately post disturbance, and decrease as stands age. Lynxes and hares utilize mid-successional stands, but are rare in young and old stands. Bats, arboreal sciurids and mustelids increase in abundance with stand age, and are most abundant in old growth. 4. Substantial gaps in the data exist for carnivores; the response of these species to fire and harvest requires research, as predator–prey interactions can affect mammal community structure in both early and late successional stages. 5. The lack of explicit treatment of in-stand forest structure post disturbance, in the reviewed literature made comparisons difficult. Where forest structure was considered, the presence of downed woody material, live residual trees and standing dead wood were shown to facilitate convergence of mammal communities to a pre-disturbance state for both disturbance types. 6. Mammalian assemblages differed considerably between successional stages, emphasizing the importance of maintaining stands of each successional stage on the landscape when implementing forest management strategies.     

Persistence of tree cavities used by cavity-nesting vertebrates declines in harvested forests

An abundant supply of cavity-bearing trees is important for maintaining wildlife communities in harvested forests. During harvesting, suitable trees and cavities are directly removed, and the longevity of cavities in retained trees may be reduced by increased exposure to wind and other disturbance factors. We examined patterns of cavity survival in retained trembling aspen (Populus tremuloides) trees in harvested stands compared with those in unharvested mature stands by monitoring the persistence of individual cavities. We followed 930 cavities in 3 harvest treatments for up to 17 years in pre-cut and uncut forest, and up to 13 years post-harvest (reserve patches and dispersed retention), in temperate-mixed forests of interior British Columbia, Canada. Average annual cavity loss rates were 5.6% in pre-cut and uncut forest, 7.2% for cavities in trees retained in reserves, and 8.1% for cavities in retained trees dispersed throughout cuts. Correspondingly, median cavity longevity was 15 years for cavities in pre-cut and uncut forest, 10 years for cavities retained in reserves, and 9 years for those in dispersed retention. Risk of loss increased most for cavities in living trees (factor of 2.17), but we found no detectable difference for cavities in recently dead trees and trees with advanced decay. We suggest retention of a range of aspen size and decay classes to allow for future cavity-tree recruitment in harvested stands. Inclusion of wildlife reserves as part of an overall forest management plan will also help to mitigate the effects of windthrow and maintain long-lived cavity resources required by a large portion of forest wildlife. © 2013 The Wildlife Society     

Stability in the patterns of long‐term development and growth of the Canadian spruce–moss forest

Aim The spruce–moss forest is the main forest ecosystem of the North American boreal forest. We used stand structure and fire data to examine the long‐term development and growth of the spruce–moss ecosystem. We evaluate the stability of the forest with time and the conditions needed for the continuing regeneration, growth and re‐establishment of black spruce (Picea mariana) trees. Location The study area occurs in Québec, Canada, and extends from 70°00′ to 72°00′ W and 47°30′ to 56°00′ N. Methods A spatial inventory of spruce–moss forest stands was performed along 34 transects. Nineteen spruce–moss forests were selected. A 500 m² quadrat at each site was used for radiocarbon and tree‐ring dating of time since last fire (TSLF). Size structure and tree regeneration in each stand were described based on diameter distribution of the dominant and co‐dominant tree species [black spruce and balsam fir (Abies balsamea)]. Results The TSLF of the studied forests ranges from 118 to 4870 cal. yr bp . Forests < 325 cal. yr bp are dominated by trees of the first post‐fire cohort and are not yet at equilibrium, whereas older forests show a reverse‐J diameter distribution typical of mature, old‐growth stands. The younger forests display faster height and radial growth‐rate patterns than the older forests, due to factors associated with long‐term forest development. Each of the stands examined established after severe fires that consumed all the soil organic material. Main conclusions Spruce–moss forests are able to self‐regenerate after fires that consume the organic layer, thus allowing seed regeneration at the soil surface. In the absence of fire the forests can remain in an equilibrium state. Once the forests mature, tree productivity eventually levels off and becomes stable. Further proof of the enduring stability of these forests, in between fire periods, lies in the ages of the stands. Stands with a TSLF of 325–4870 cal. yr bp all exhibited the same stand structure, tree growth rates and species characteristics. In the absence of fire, the spruce–moss forests are able to maintain themselves for thousands of years with no apparent degradation or change in forest type.     

Sources of variation in the incidence of ant–aphid mutualism in boreal forests

• 1 The mutualism between wood ants of the Formica rufa group and aphids living in the canopy of trees is a widespread phenomenon in boreal forests, and it can affect tree growth. However, not all trees in the forest are involved in this interaction.
• 2 To assess the incidence of host trees involved in this ant–aphid mutualism and its spatial distribution in boreal forests, we inventoried sample plots with a radius of 10–15 m around wood ant mounds in 12 forest stands of two age classes (5–12‐year‐old sapling stands and 30–45‐year‐old pole stands) and two dominant tree species (Scots pine and silver birch) in Eastern Finland from 2007 to 2009.
• 3 The proportion of trees visited by ants out of all trees on the individual study plots were in the range 4–62%, and 1.5–39% of the trees on the plots were consistently visited by ants during all 3 years. The percentage of host trees increased with the ant mound base area on the plots. Trees visited by ants were larger and closer to the mound than trees not visited by ants. Within the group of visited trees, more ants were found on bigger trees and on trees close to the ant mounds.
• 4 Extrapolated from plot to stand level, we estimated that 0.5–6.6% of the trees were host trees in at least one of the three study years, and that only 0.01–2.3% of all the trees were consistently visited by ants during all 3 years. It is concluded that ant–aphid mutualism is a minor occurrence at the stand level.

     

Understory vegetation response to thinning disturbance of varying complexity in coniferous stands

Question: Can augmented forest stand complexity increase understory vegetation richness and cover and accelerate the development of late‐successional features? Does within‐stand understory vegetation variability increase after imposing treatments that increase stand structural complexity of the overstory? What is the relative contribution of individual stand structural components (i.e. forest matrix, gaps, and leave island reserves) to changes in understory vegetation richness? Location: Seven study sites in the Coastal Range and Cascades regions of Oregon, USA. Methods: We examined the effects of thinning six years after harvest on understory plant vascular richness and cover in 40‐ to 60‐year‐old forest stands dominated by Douglas‐fir (Pseudotsuga menziesii). At each site, one unthinned control was preserved and three thinning treatments were implemented: low complexity (LC, 300 trees ha^?1), moderate complexity (MC, 200 trees ha^?1), and high complexity (HC, variable densities from 100 to 300 trees ha^?1). Gaps openings and leave island reserves were established in MC and HC. Results: Richness of all herbs, forest herbs, early seral herbs and shrubs, and introduced species increased in all thinning treatments, although early seral herbs and introduced species remained a small component. Only cover of early seral herbs and shrubs increased in all thinning treatments whereas forest shrub cover increased in MC and HC. In the understory, we found 284 vascular plant species. After accounting for site‐level differences, the richness of understory communities in thinned stands differed from those in control stands. Within‐treatment variability of herb and shrub richness was reduced by thinning. Matrix areas and gap openings in thinned treatments appeared to contribute to the recruitment of early seral herbs and shrubs. Conclusions: Understory vegetation richness increased 6 years after imposing treatments, with increasing stand complexity mainly because of the recruitment of early seral and forest herbs, and both low and tall shrubs. Changes in stand density did not likely lead to competitive species exclusion. The abundance of potentially invasive introduced species was much lower compared to other plant groups. Post‐thinning reductions in within‐treatment variability was caused by greater abundance of early seral herbs and shrubs in thinned stands compared with the control. Gaps and low‐density forest matrix areas created as part of spatially variably thinning had greater overall species richness. Increased overstory variability encouraged development of multiple layers of understory vegetation.     

Influence of canopy tree size on stand basal area may reflect uncoupling of crown expansion and trunk diameter growth

Abstract A recent article by Midgley and colleagues suggests that large trees give rise to inordinately high stand basal areas because they pack canopy space more efficiently than smaller trees. We argue that this phenomenon bears more relation to the fact that diameter increment is not necessarily accompanied by significant crown expansion during all stages of a tree's life. Using data from a canopy tree population in an old‐growth temperate forest, we found that crown area scaled as roughly the 3/5 power of trunk basal area. Rather than reflecting fixed scaling laws, we suggest that this pattern arises because of limited opportunities for crown expansion in dense stands. Old canopy trees in dense stands can thus accumulate large basal areas without occupying a commensurately large canopy area.     

Radial and stand‐level thinning treatments: 15‐year growth response of legacy ponderosa and Jeffrey pine trees

Restoration efforts to improve vigor of large, old trees and decrease risk to high‐intensity wildland fire and drought‐mediated insect mortality often include reductions in stand density. We examined 15‐year growth response of old ponderosa pine (Pinus ponderosa) and Jeffrey pine (Pinus jeffreyi) trees in northeastern California, U.S.A. to two levels of thinning treatments compared to an untreated (control) area. Density reductions involved radial thinning (thinning 9.1 m around individual trees) and stand thinning. Annual tree growth in the stand thinning increased immediately following treatment and was sustained over the 15 years. In contrast, radial thinning did not increase growth, but slowed decline compared to control trees. Available soil moisture was higher in the stand thinning than the control for 5 years post‐treatment and likely extended seasonal tree growth. Our results show that large, old trees can respond to restoration thinning treatments, but that the level of thinning impacts this response. Stand thinning must be sufficiently intensive to improve old tree growth and health, in part due to increasing available soil moisture. Importantly, focusing stand density reductions around the immediate neighborhood of legacy trees was insufficient to elicit a growth response, calling into question treatments attempting to increase vigor of legacy trees while still maintaining closed canopies in dry, coniferous forest types. Although radial thinning did not affect tree growth rates, this treatment may still achieve other resource objectives not studied here, such as protecting wildlife habitat, reducing the risk of severe fire injury, and decreasing susceptibility to bark beetle attacks.     

Carbon and nitrogen storage in an age-sequence of <Emphasis Type="Italic">Pinus densiflora</Emphasis> stands in Korea

The carbon (C) and nitrogen (N) storage capabilities of Pinus densiflora in six different stand ages (10, 27, 30, 32, 44, and 71 years old) were investigated in Korea. Thirty sample trees were destructively harvested and 12 were excavated. Samples from the above and belowground tree components, coarse woody debris (CWD), forest floor, and mineral soil (0–30 cm) were collected. Tree biomass was highest in the 71-year-old stand (202.8 t ha⁻¹) and lowest in the 10-year-old stand (18.4 t ha⁻¹). C and N storage in the mineral soil was higher in the 71-year-old stand than in the other stands, mainly due to higher soil C and N concentrations. Consequently, the total ecosystem C and N storage (tree+forest floor+CWD+soil) was positively correlated with stand age: increasing from a minimum in the 10 year old stand (18.8 t C ha⁻¹ and 1.3 t N ha⁻¹) to a maximum in the 71-year-old stand (201.4 t C ha⁻¹ and 8.5 t N ha⁻¹). The total ecosystem C storage showed a similar sigmoidal pattern to that of tree C storage as a function of the age-sequence, while N storage in the CWD, forest floor and mineral soil showed no significant temporal trends. Our results provide important insights that will increase our understanding of C and N storage in P. densiflora stands and our ability to predict changes according to stand age in the region.     

Carbon isotope discrimination and growth response of old Pinus ponderosa trees to stand density reductions

Stand density reductions have been proposed as a method by which old‐growth ponderosa pine (Pinus ponderosa) forests of North America can be converted back to pre‐1900 conditions, thereby reducing the danger of catastrophic forest fires and insect attacks while increasing the productivity of the remaining old‐growth individuals. However, the duration of productivity response of individual trees and the physiological mechanisms underlying such a response remain speculative issues, particularly in old trees. Tree‐ring measurements of carbon isotope ratios (δ¹³C) and basal area increment (BAI) were used to assess the response of intrinsic water‐use efficiency (the ratio of photosynthesis, A to stomatal conductance, g) and growth of individual> 250‐year‐old‐ponderosa pine trees to stand density reductions. It was hypothesized that reductions in stand density would increase soil moisture availability, thus decreasing canopy A/g and increasing carbon isotope discrimination (Δ). Cellulose‐δ¹³C of annual tree rings, soil water availability (estimated from pre‐dawn leaf water potential), photosynthetic capacity, stem basal growth and xylem anatomy were measured in individual trees within three pairs of thinned and un‐thinned stands. The thinned stands were treated 7 to 15 years prior to measurement. The values of δ¹³C and BAI were assessed for 20 consecutive years overlapping the date of thinning in a single intensively studied stand, and was measured for 3 years on either side of the date of thinning for the two other stands to assess the generality of the response. After thinning, Δ increased by 0.89‰ (± 0.15‰). The trees in the un‐thinned stands showed no change in Δ (0.00‰ ± 0.04‰). In the intensively studied trees, significant differences were expressed in the first growing season after the thinning took place but it took 6 years before the full 0.89‰ difference was observed. BAI doubled or tripled after disturbance, depending on the stand, and the increased BAI lasted up to 15 years after thinning. In the intensively studied trees, the BAI response did not begin until 3 years after the Δ response, peaked 1 year after the Δ peak, and then BAI and Δ oscillated in unison. The lag between BAI and Δ was not due to slow changes in anatomical properties of the sapwood, because tracheid dimensions and sapwood‐specific conductivity remained unchanged after disturbance. The Δ response of thinned trees indicated that A/g decreased after thinning. Photosynthetic capacity, as indexed by foliar nitrogen ([N]) and by the relationship between photosynthesis and internal CO₂ (A–C_i curves), was unchanged by thinning, confirming our suspicion that the decline in A/g was due to a relatively greater increase in g in comparison with A. Model estimates agreed with this conclusion, predicting that g increased by nearly 25% after thinning relative to a 15% increase in A. Pre‐dawn leaf water potential averaged 0.11 MPa (± 0.03 MPa) less negative for the thinned compared with the un‐thinned trees in all stands, and was strongly correlated with Δ post‐thinning (R² = 0.91). There was a strong relationship between BAI and modelled A, suggesting that changes in water availability and g have a significant effect on carbon assimilation and growth of these old trees. These results confirm that stand density reductions result in increased growth of individual trees via increased stomatal conductance. Furthermore, they show that a physiological response to stand density reductions can last for up to 15 years in old ponderosa pines if stand leaf area is not fully re‐established.     

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.