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

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

Analysis and modeling of spatial stand structures. Methodological considerations based on mixed beech-larch stands in Lower Saxony

The first part of this paper highlights spatial stand structure as the central stand characteristic and introduces methods of pattern identification. This involves two nearest-neighbour methods for the identification of stand structures, i.e., the aggregation index R by Clark and Evans [Clark, Ph.J. and Evans, F.C., 1954. Distance to nearest neighbor as a measure of spatial relationships in populations. Ecology 35 (4) 445–453.] for univariate patterns and the segregation index S by Pielou [Pielou, E.C., 1977. Mathematical Ecology. Wiley.] for bivariate patterns. Both were used to describe the structure of 53 experimental plots of mixed beech-larch stands in Lower Saxony which provided the data base for this investigation. The second part of the study deals with the development of the STRUGEN stand structure generator, designed for the modeling and reproduction of spatial stand structures. To generate stand structures, a two-dimensional homogeneous Poisson process is used as well as a set of two-dimensional distribution functions which determine mixture type and intermingling intensity of main and associated tree species. Moreover, a distance function secures minimum distances between competing neighbouring trees. Consequently, the produced pattern is the result of a combination of an inhomogeneous Poisson process (for generating mixture units) and a hard-core process (for securing minimum distances between neighbours). The STRUGEN generator was designed and successfully used for the investigation of 53 mixed beech-larch stands. It provides initial values and stand structures for distance-dependent single-tree models from estimated qualitative stand characteristics. STRUGEN is a useful tool and allows initial, pragmatic steps towards fully utilising available qualitative and quantitative information to diagnose the state of a forest and to predict its growth.     

林分空间结构参数角尺度的标准角选择

角尺度是一个刻画森林空间结构的新参数,通过描述相邻木围绕参照树的均匀性来进行林木分布格局的判定。角尺度对复杂森林的空间结构有很强的解析能力。标准角是衡量参照树周围4株最近相邻木分布均匀性的标准,是影响角尺度使用精度的关键因子。理论推导出的标准角应为72°,这个标准角能使随机分布林分的角尺度均值最接近0.5,从而与角尺度定义中林木随机分布时角尺度取值为0.5相一致,这个观察被进一步模拟的100个随机分布的林分所证实。     

基于目标树经营的抚育采伐对云冷杉针阔混交林空间结构的影响

[目的]以长白山林区云冷杉针阔混交林为研究对象,分析目标树抚育采伐前后云冷杉针阔混交林空间结构参数的变化,探索目标树抚育采伐对森林空间结构的影响。[方法]基于6块面积1 hm~2的样地,首先遵循近自然森林经营的原理进行林木分类,选定目标树,伐除干扰树;再结合结构化森林经营的原理,计算并分析抚育采伐前后目标树空间结构参数和林分空间结构评价指数的变化。[结果]表明:经目标树抚育采伐后(1)目标树空间结构单元内的树种隔离程度得到进一步提高;林木透光条件有所改善;目标树的竞争压力得以减小;林木空间分布格局总体上得到优化。(2)林分空间结构评价指数进一步提高,林分空间结构水平稳中有升。[结论]通过采取合理的目标树抚育采伐措施能兼顾改善单木水平和林分水平的空间结构状况,在实现优化目标树空间结构的同时,保证林分整体空间结构的稳定。从目标树抚育采伐对林分整体空间结构的影响来看:所有样地的林分空间结构状况均得到进一步改善。     

Windthrow probability as a function of stand characteristics and shelter

In 1981 a storm caused windthrow of 3 million cubic meters of softwood in Denmark, equal to the normal removals of three years. The object of the present analysis is to determine the way in which the probability of windthrow depends on parameters that can be affected by forest management, viz. rotation age, thinning programme, choice of species, spatial distribution of stands and drainage. An empirical data set from 612 stands is used in the estimations. It is found that the windthrow probability is negatively affected by tree diameter, drainage, the time since last thinning and the protection from other stands. The probability increases with tree height, age and relative thinning volume in the latest thinning. Picea is more stable than Abies and Pseudotsuga.     

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

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

     

Canopy vs. understory: Does tree diversity affect bee and wasp communities and their natural enemies across forest strata?

Bees and wasps provide important ecosystem services such as pollination and biocontrol in crop-dominated landscapes, but surprisingly little information is available on hymenopteran communities in temperate forest ecosystems. Species richness and abundance of bees and wasps can be hypothesised to increase with plant diversity, structural complexity, and availability of food and nesting resources. By experimentally exposing standardised nesting sites, we examined abundance and species richness of cavity-nesting bees (pollinators), wasps (predators) and their associated parasitoids across a tree diversity gradient in a temperate deciduous forest habitat. In addition, spatial distribution of individuals and species across forest strata (canopy vs. understory) was tested. Abundance and species richness was high for predatory wasps, but generally low for pollinators. Species-rich forest stands supported increased abundance, but not species richness, of pollinators and predatory wasps, and also increased abundance and species richness of natural enemies. In addition, the forests showed a distinct spatial stratification in that abundance of bees, wasps and parasitoids as well as parasitism rates were higher in the canopy than understory. We conclude that particularly the canopy in temperate forest stands can serve as an important habitat for predatory wasp species and natural enemies, but not bee pollinators. Enhanced tree diversity was related to increased hymenopteran abundance, which is likely to be linked to an increase in nesting and food resources in mixed forest stands.     

Nature conservation and timber production in areas with fragmented ownership patterns

Forestry has transformed the tree species composition and structure of Swedish forests. The fragmented ownership pattern in areas with non-industrial private forest ownership (NIPF), in combination with these forestry practices, have created fragmented forests with relatively low proportions of habitat types important to many species. Ecological landscape planning has been suggested and tested as a mean for integrating nature conservation and timber production in Sweden. However, ecological landscape planning concepts have been developed for areas with homogenous ownership patterns and not for areas with fragmented ownership. In this study, stands that are voluntarily set aside by individual forest owners were examined in terms of nature conservation value, and compared to randomly selected stands obtained by Monte Carlo simulation. In order to obtain rather detailed data about the set-aside stands, semi-standardized interviews were carried out directly with forest owners in three landscape areas dominated by NIPF. The interviews showed that 26 out of 29 forest owners could locate some stands where they did not have high requirements for timber yield. These stands had higher than average local and spatial nature conservation value as a result of stand structure, composition and location in the landscape. The stands did not have a random location or size, being found closer to key habitats as well as forming equally large or larger patches than other stands. These stands are suggested to form a basis for ecological landscape planning as forest owners seem to have some kind of ‘common view’ of which stands to set aside. This common view could be used to coordinate the contribution made by individual forest owners to nature conservation in areas with fragmented ownership patterns.     

Promoting old-growth characteristics and long-term wood production in Douglas-fir forests

Trade-offs among wood production, wood quality and ecological characteristics in the management of harvested forest stands are explored through model simulation of various silvicultural regimes. Long-term production of merchantable wood, production of various types of high-quality wood, and the level of certain quantitative ecological indicators are projected for coniferous forests of Pacific Northwestern USA. The set of ecological indicators used is based on the species composition and physical structure of old, unlogged forest stands. Simulations are performed with an ecological model of forest stand dynamics that tracks the fate of live and dead trees. Short rotations (<50 years) produce the least amount of high-quality wood over the multi-century simulation period. They also fail to generate ecological attributes resembling those of old forest stands. Production of high-quality wood is moderate to high under all rotations of 80 years or more; however, most ecological indicators require longer rotations unless alternatives to clearcutting are applied. Alternatives examined include retention of 15% cover of live tree canopy at each harvest in combination with artificial thinning between harvests. Thinning from below can expedite the development of large live and dead trees, and canopy height diversity without greatly diminishing wood quantity or quality. Proportional thinning retains understory stems, thereby expediting the recruitment of shade-tolerant trees. A possible drawback to thinning, particularly proportional thinning, is the diminished production of clean-bole wood at rotations of 150 and 260 years. It is concluded that most wood quantity, wood quality and ecological objectives can be met with long rotations (ca. 260 years). Certain objectives can be met with shorter rotations (80–150 years) when treatments of thinning and canopy tree retention are applied.     

修枝间伐对松突圆蚧抑制作用的研究

严重危害广东省沿海地区马尾松的松突圆蚧(Hemiberlesia pitysophila Takagi),自1982年在深圳、珠海等地发现后，至今危害面积约43万ha，其中枯萎或涉死的约8万ha。     

Effects of thinning and canopy type on growth dynamics of Pinus sylvestris: inter-annual variations and intra-annual interactions with microclimate

We assessed the effects of thinning (0, 20 and 30 % extraction of basal area) and canopy type (pine–beech vs. pine plots, beech accounting for 12 % of total basal area) on radial growth of dominant and codominant Scots pine at inter-annual scale and on microclimatic conditions, radial growth and xylogenesis 9 years after thinning at intra-annual scale. Thinning weakly affected pine growth, which was enhanced 3 years after harvesting. Over time, a gradual reduction in pine growth in mixed canopy relative to pure canopy occurred only in unthinned plots apparently due to beech expansion. Indeed, 9 years after thinning, a higher seasonal radial increment and a greater number of tracheids were produced under pine canopy in the unthinned plots, whereas no differences between canopy types were observed in the thinned plots. Radial increment and tracheid production were mainly affected by tree water status (air and soil humidity, throughfall). The differences of tree water status caused by treatments, and plausibly disparities in tree size and tree-to-tree competition, were the main drivers explaining the patterns observed for radial increment and xylogenesis. Our results suggest that the negative effects of beech competition on Scots pine growth in similar mixed forest may be controlled to some extent by thinning.     

Model-based investigation on the effects of spatial evenness,and size selection in thinning of Picea abies stands

Size and spatial distribution of trees are important for forest stand growth, but the extent to which it matters in thinning operations, in terms of wood production and stand economy, has rarely been documented. Here we investigate how the choice of spatial evenness and tree-size distribution of residual trees impacts wood production and stand economy. A spatially explicit individual-based growth model was used, in conjunction with empirical cost functions for harvesting and forwarding, to calculate net production and net present value for different thinning operations in Norway spruce stands in Northern Sweden. The in silico thinning operations were defined by three variables: (1) spatial evenness after thinning, (2) tree size preference for harvesting, and (3) basal area reduction. We found that thinning that increases spatial evenness increases net production and net present value by around 2.0%, compared to the worst case. When changing the spatial evenness in conjunction with size preference we could observe an improvement of the net production and net present value up to 8.0%. The magnitude of impact differed greatly between the stands (from 1.7% to 8.0%) and was highest in the stand with the lowest stem density.     

Spatially explicit assessment of carbon stocks of a managed forest area in eastern Germany

The Kyoto-protocol permits the accounting of changes in forest carbon stocks due to forestry. Therefore, forest owners are interested in a reproducible quantification of carbon stocks at the level of forest management units and the impact of management to these stocks or their changes. We calculated the carbon stocks in tree biomass and the organic layer including their uncertainties for several forest management units (Tharandt forest, Eastern Germany, 5,500 ha) spatially explicit at the scale of individual stands by using standard forest data sources. Additionally, soil carbon stocks along a catena were quantified. Finally, carbon stocks of spruce and beech dominated stands were compared and effects of thinning intensity and site conditions were assessed. We combined forest inventory and data of site conditions by using the spatial unions of the shapes (i.e., polygons) in the stand map and the site map. Area weighted means of carbon (C) stocks reached 10.0 kg/m2 in tree biomass, 3.0 kg/m2 in the organic layer and 7.3 kg/m2 in mineral soil. Spatially explicit error propagation yielded a precision of the relative error of carbon stocks at the total studied area of 1% for tree biomass, 45% for the organic layer, and 20% for mineral soil. Mature beech dominated stands at the Tharandt forest had higher tree biomass carbon stocks (13.4 kg/m2) and lower organic layer carbon stocks (1.8 kg/m2) compared to stands dominated by spruce (11.6, 3.0 kg/m2). The difference of tree biomass stocks was mainly due to differences in thinning intensity. The additional effect of site conditions on tree carbon stocks was very small. We conclude that the spatially explicit combination of stand scale inventory data with data on site conditions is suited to quantify carbon stocks in tree biomass and organic layer at operational scale.     

采伐对天然云冷杉针阔混交林半分解层凋落物现存量、含水率及林分郁闭度空间异质性的影响

[目的]以吉林省汪清林业局金沟岭林场4块天然云冷杉针阔混交林为研究对象,探究不同采伐强度下[重度(采伐强度21. 21%),中度(采伐强度11. 22%),轻度(采伐强度6. 29%)和对照(未采伐)]凋落物半分解层现存量、含水率与林分郁闭度的空间异质性。[方法]采用地统计学方法,对各样地凋落物半分解层现存量、含水率与林分郁闭度数据进行普通克里金插值,得到3个指标的空间分布格局图。[结果]表明:在10 100 m的尺度上,采伐使得凋落物现存量由离散型分布转变为聚集型分布,而对凋落物含水率的空间格局影响不大;所有样地的郁闭度均表现为聚集型分布,采伐一定程度上影响了其在中度采伐与重度采伐样地的空间异质性;凋落物现存量与林分郁闭度之间不存在相关性,而在对照样地,凋落物现存量与含水率存在显著的负相关,但相关程度较弱(r 0. 3)。[结论]采伐对凋落物半分解层含水率空间异质性影响不大;中度采伐与重度采伐样地的郁闭度异质性可能是由于采伐不均匀所导致;采伐使凋落物现存量空间异质性程度降低。     

Spatial point-pattern analysis for detecting density-dependent competition in a boreal chronosequence of Alberta

In the boreal forest of Alberta, fire and wind often open gaps in the canopy where late-successional species can establish and over time cause a shift in the species distribution from deciduous (e.g., trembling aspen) dominated to mixedwood, to shade-tolerant conifer (e.g., white spruce) dominated stands. This study attempted to understand the change of density-dependent competition in a boreal chronosequence and the role of tree competition in affecting stand structure and mortality. Four 1-ha stem-mapped plots were established to represent a chronosequence comprised of aspen dominated, mixedwood, and spruce dominated stands in Alberta. Second order spatial point-pattern analysis using Ripley's K(t) function showed that intraspecific competition is a prevailing force causing conspecific tree mortality and thus shaping the stand structure. The results of bivariate K(t) function analysis did not reveal sufficient evidence of interspecific competition. This suggested that competitive interaction among heterospecific trees was not strong enough to cause significant tree mortality, but the analysis of marked correlation function revealed that interspecific competition could have a negative impact on tree growth. This study highlights the importance of density-dependent competition in understanding stand dynamics of boreal forests over succession.     

Review of ground-based methods to measure the distribution of biomass in forest canopies

? Context

Ecological research and an effective forest management need accurate information on the structure of the forest canopy to understand the biochemical, physiological and biogeochemical processes within a forest.

? Research question

This paper reviews the currently available instruments for measuring the distribution of biomass within forest canopies. We compare the most well-established approaches and present the different measurable parameters. A special focus lies on the resolution of the obtained data.

? Results

It was found that only 3D laser scanners offer data with the resolution required by ecologists, private landholders, the forest industry and the public to detect trends in tree growth patterns and canopy interactions in all three spatial dimensions. But data validation, data analysis and parameter extraction are still under development, and the price of the instrument is quite high.

? Conclusion

Research should focus on the parameter extraction from terrestrial laser scanner data as this could allow the calculation of functional attributes for different sections of a canopy on a high spatial resolution. It could also help ecologists characterize the structure of forest stands in a quick and precise way.     

Canopy light transmittance in Douglas-fir--western hemlock stands

We measured vertical and horizontal variation in canopy transmittance of photosynthetically active radiation in five Pseudotsuga menziesii (Mirb.) Franco-Tsuga heterophylla (Raf.) Sarg. (Douglas-fir-western hemlock) stands in the central Cascades of southern Washington to determine how stand structure and age affect the forest light environment. The shape of the mean transmittance profile was related to stand height, but height of mean maximum transmittance was progressively lower than maximum tree height in older stands. The vertical rate of attenuation declined with stand age in both the overstory and understory. A classification of vertical light zones based on the mean and variance of transmittance showed a progressive widening of the bright (low variance and high mean) and transition (high variance and rapid vertical change) zones in older stands, whereas the dim zone (low variance and mean) narrowed. The zone of maximum canopy surface area in height profiles, estimated by inversion of transmittance profiles, changed from relatively high in the canopy in most young stands ("top-heavy") to lower in the canopy in older stands ("bottom-heavy"). In the understory, all stands had similar mean transmittances, but the spatial scale of variation increased with stand age and increasing crown size. The angular distribution of openness was similar in all stands, though the older stands were less open at all angles than the younger stands. Understory openness was generally unrelated to transmittance in the canopy above. Whole-canopy leaf area indices, estimated using three methods of inverting light measurements, showed little correspondence across methods. The observed patterns in light environment are consistent with structural changes occurring during stand development, particularly the diversification of crowns, the creation of openings of various sizes and the elaboration of the outer canopy surface. The ensemble of measurements has potential use in distinguishing forests of differing ages that have similar stature.     

Altered light conditions following thinning affect xylem structure and potential hydraulic conductivity of Norway spruce shoots

Trees must respond to many environmental factors during their development, and light is one of the main stimuli regulating tree growth. Thinning of forest stands by selective tree removal is a common tool in forest management that increases light intensity. However, morphological and anatomical adaptations of individual shoots to the new environmental conditions created by thinning are still poorly understood. In this study, we evaluated shoot morphology (shoot length, needle number, projected leaf area) and anatomy (tracheid lumen area, tracheid number, tracheid dimensions, xylem area, potential hydraulic conductivity) in three Norway spruce (Picea abies/L./Karst.) families exposed to different thinning regimes. We compared shoot characteristics of upper-canopy (i.e. sun-exposed) and lower-canopy (i.e. shaded) current-year shoots in a control plot and a plot thinned to 50 % stand density the previous year. One tree per family was chosen in each treatment, and five shoots were taken per canopy position. We found that upper-canopy shoots in both plots had higher values than lower-canopy shoots for all studied parameters, except lumen roundness and tracheid frequency (i.e. tracheid number per xylem area). Thinning had little effect on shoot morphology and anatomy 1 year after thinning, except for small but significant changes in tracheid dimensions. Needles were more sensitive to altered light conditions, as projected leaf area of shoot, needle number and leaf hydraulic conductivity changed after thinning. Differences between upper- and lower-canopy shoots did not seem to be influenced by thinning and were almost the same in both plots. Our results suggest that lower-canopy shoots require several years to modify their morphology and anatomy to new light conditions following thinning. The slow light adaptation of the lower canopy may be of practical importance in forest management: thinned stands may be predisposed to drought stress because newly exposed shoots experience increased illumination and transpiration after thinning.     

A preliminary study on the effects of line and selective thinning on bird communities in Hokkaido, northern Japan

In Japan, selective thinning is a common thinning method, though line thinning receives much attention because of its economic merits. In this study, we examined effects of the two thinning methods on bird communities in Todo fir (Abies sachalinensis) plantations in Hokkaido, Japan. We surveyed bird species in forests under four different management types — unthinned, selectively thinned, line-thinned plantation, and naturally regenerated forest (here after referred to as natural forest) stands — using a line-transect method. We also investigated vegetation structure (canopy tree and understory) of these stands. Bird species richness did not differ between natural forests and plantations, while bird total abundance was greater in plantations than in natural forests. Bird species richness and total abundance were comparable among the three management types for plantations. Abundances of 10 bird species were different among the four management types, and five species were more abundant in line-thinned plantations. However, two species were more abundant in selectively thinned stands than in line-thinned stands, and they frequently appeared in natural forests. There were no distinct differences in vegetation structure among the management types for plantations. Our results suggest that line thinning could be beneficial for some bird species in plantations.     

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

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