Variation in post-wildfire regeneration of boreal mixedwood forests: underlying factors and implications for natural disturbance-based management

To test the direct regeneration hypothesis and support natural disturbance-based forest management we characterized the structure and composition of boreal mixedwood forests regenerating after large wildfires and examined the influence of pre-fire stand composition and post-fire competing vegetation. In stands which had been deciduous (Populus sp.)-dominated, conifer (white spruce)-dominated, or mixed pre-fire we measured regeneration stocking (presence in 10 m² plots), density and height 10–20 years post-burn in five wildfires in Alberta, Canada. Most plots regenerated to the deciduous or mixed stocking types; plots in the older fire and in stands that were pure conifer pre-fire had higher amounts of conifer regeneration. Surprisingly, regeneration in pre-fire ‘pure’ white spruce stands was most often to pine, although these had not been recorded in the pre-fire inventory. Pre-fire deciduous stands were the most resilient in that poplar species dominated their post-fire regeneration in terms of stocking, density and height. These stands also had the highest diversity of regenerating tree species and the most unstocked plots. High grass cover negatively affected regeneration density of both deciduous and conifer trees. Our results demonstrate the natural occurrence of regeneration gaps, pre- to post-fire changes in forest composition, and high variation in post-fire regeneration composition. These should be taken into consideration when developing goals for post-harvest regeneration mimicking natural disturbance.     

Comparison of relascope and fixed-radius plots for the estimation of forest stand variables in northeast Spain: an inventory simulation approach

The Bitterlich relascope is a multiple use dendrometer widely used in forest inventory. Although it is most commonly used to estimate basal area, the relascope can also estimate other stand variables, including density and diameter distribution. However, forest stand inventories in Spain rarely use relascope plots to estimate these variables due to the belief that they lead to higher errors than fixed-radius plots due to the heterogeneity of many Mediterranean forests. This study compared the accuracy of the estimated averages of three main stand variables (basal area, stand density, and diameter class distribution) in forest stand inventories performed with relascope plots and with conventional fixed-radius circular plots, both measuring a similar number of trees (15–20). A forest stand inventory simulator (DOMO) was used (1) to generate simulated forest stands corresponding to the nine most common types in the Mediterranean region of Catalonia (NE Spain), including even-aged and uneven-aged stands, and (2) to estimate and compare the average values of these variables at the forest stand level resulting from both plot types. In general, we did not find significant accuracy differences between the inventory systems for most of the stand variables and forest types studied, as expected by established angle-count sampling theory. However, the results show that for stands with multiple strata and open structures, the Bitterlich relascope provides a more accurate estimate for basal area than for density, while the reverse occurs for fixed-radius plots.     

Water status and drought stress in experimental gaps in managed and semi-natural silver fir--beech forests

The effect of gap characteristics, including gap age, vegetation cover, and stage of regeneration, on water balance and drought stress was studied over a four-year period in natural and experimental gaps of various sizes in Dinaric silver fir--beech forest in Slovenia. A medium gap, two experimental clear-cut gaps, and a closed stand were selected in a traditional irregular shelterwood managed forest, and an irregular-shaped natural gap and part of the stand were selected in a semi-natural forest. Soil moisture, throughfall, and stemflow were measured in the gaps and stands, and incident precipitation was monitored in the open area. Evapotranspiration and drainage water fluxes were estimated using the water balance model BROOK90. To estimate drought stress, the ratio between simulated actual and potential evapotranspiration was used. Precipitation in the 2001 and 2003 growing seasons was considerably less than that in 2002 and 2004. The highest drainage fluxes were in the newly created clear-cut gaps with sparse ground vegetation cover, followed by the natural gap and medium gap, which had older and denser natural tree regeneration. On average, the clear-cut gap drainage fluxes were 18% (164?mm) higher than those in the forest stands. Evapotranspiration was lower in the natural gap and higher in the managed gap compared with the experimental clear-cut gaps. Water supply stress was greater for the forest stands than for the vegetation in the gaps. Drought stress was indicated to be lowest in the natural gap, which had patches of varying development phases and the highest water storage capacity of the soil and vegetation.     

Planning timber harvest of residual forest stands without compromising bird and small mammal communities in boreal landscapes

Numerous efforts have been invested in designing and configuring residual forest stands in Canadian boreal forest to preserve their overall biodiversity. Now that several landscapes have been partially logged, the next issue in forest management involves the planning of residual forest stand harvesting without compromising wildlife populations. Residual stands can be cut when adjacent regeneration reaches 3 m in height according to current regulations in several Canadian provinces (e.g., Québec, Ontario, Alberta, and British-Columbia). However, little is known on whether such regenerating habitat (RE-3m) can maintain wildlife communities similar to those found in unharvested mature forest (CO). We estimated the relative abundance of small mammals and forest birds in RE-3m and CO habitats and characterized landscape and stand structures. These variables were then compared between the two contrasting successional stages and were used to build habitat use models (HUMs) for 21 species. CO and RE-3m differed with regard to several landscape characteristics and stand structure variables as a result of logging. Snowshoe Hare, Northern Flicker, Alder Flycatcher, Ruby-crowned Kinglet, White-throated Sparrow and Magnolia Warbler were more abundant in RE-3m than CO, while Red-backed Vole, Brown Creeper and Golden-crowned Kinglet exhibited lower abundances in RE-3m. No significant differences in abundance were observed for the 12 other species. Species HUMs were highly significant and explained between 64.3 and 99.1% of the total variability in abundance. Following variance partitioning, stand structure variables accounted for most of the explained variability (54.2%) while landscape characteristics accounted for only 28.7%. No difference in species richness was observed but community evenness was greater in CO than RE-3m. Our results suggest that current regulations may threaten the maintenance of 3 out of 21 censused species for which abundances were significantly lower in regenerating 3 m tall stands. As stand structure explained a large amount of variability in abundance, it should be considered during timber harvest planning in both mature and regenerating stands. Until we know more on whether the current regulations are suitable for maintaining overall biodiversity, our results suggest that some mature forest stands should be maintained within managed landscapes for a complete logging rotation period.     

Do changes in spatial distribution, structure and abundance of silver fir (Abies alba Mill.) indicate its decline?

Silver fir decline was investigated based on changes in spatial distribution of fir, fir abundance in forest stands, dbh (age) structure of fir, and abundance of fir regeneration. The authors used a large-scale approach to study the dynamics of silver fir over nearly 40 years. Based on Silva-SI, a spatial information system, the majority of total forest area in Slovenia was analysed for changes in the distribution of silver fir in the period 1970-2008 using artificial neural networks (ANNs), with respect to site, stand, and forest management variables. Additionally, changes in abundance of silver fir (hereafter fir) in the same period were analysed with a general linear model, while changes in the dbh structure of fir and fir regeneration were analysed with non-parametric tests. Most selected indicators confirmed the hypothesis of fir decline in the period 1970-2008, as evidenced by a reduction in the area of forests with a share of fir in the total growing stock >25% (from 18.9% to 9.5% of total area), a reduction in the share of fir in the growing stock of forest stands (from 17.5% to 7.5%), ageing of the fir population, and a disproportionate share of fir saplings in the total saplings relative to fir's share in the growing stock of forest stands. A 1.5% increase in fir distribution area in the observed period contradicts the decline hypothesis. ANNs showed that the spatiotemporal dynamics of fir was most affected by four variables: forest type, share of fir in the potential natural vegetation, mean annual precipitation, and mean annual temperature. The latter two, together with the growing stock at the start of study period, the degree of connectivity between fir stands, and bedrock type, were significant predictors of decline of fir abundance in forest stands. Significant differences in spatiotemporal dynamics and changes in fir abundance were found between forest types representing a complex of site conditions and past forest management. A further decline of abundance of fir on a large spatial scale in the next decades is expected. Conservation of fir in forest stands will depend mainly on the successful regeneration of fir and sufficient recruitment.     

Structure of a Central-European mountain spruce old-growth forest with respect to historical development

This study examines the structural characteristics of the tree layer, dead wood, canopy openings, and regeneration patterns of a spruce old-growth forest in the Bohemian Forest, Czech Republic. An old-growth stand with minor human influence and a stand that was presumably logged about 200 years ago were analyzed and compared, as some forest managers considered the presumable human impact as a reason for salvage logging. Even though the stands differed in tree density, height and DBH structure, it was not possible to conclude whether it was due to management history or the environmental differences. The volume of dead wood also differed between the stands. There was about 142 and 83 m³ ha⁻¹ of dead wood in the old-growth stand and presumably logged stand, respectively. The amount of dead wood found in the old-growth stand was comparable with values reported from spruce old-growth stands across Central Europe. In both stands, many canopy trees were arranged in linear patterns, which was a result of spruce regeneration on nurse logs. This suggests that the origin and development of the stands were characterized by natural processes and during the past 200 years typical old-growth structural characteristics have already evolved.     

A silviculture-oriented spatio-temporal model for germination in Pinus pinea L. in the Spanish Northern Plateau based on a direct seeding experiment

Natural regeneration in Pinus pinea stands commonly fails throughout the Spanish Northern Plateau under current intensive regeneration treatments. As a result, extensive direct seeding is commonly conducted to guarantee regeneration occurrence. In a period of rationalization of the resources devoted to forest management, this kind of techniques may become unaffordable. Given that the climatic and stand factors driving germination remain unknown, tools are required to understand the process and temper the use of direct seeding. In this study, the spatio-temporal pattern of germination of P. pinea was modelled with those purposes. The resulting findings will allow us to (1) determine the main ecological variables involved in germination in the species and (2) infer adequate silvicultural alternatives. The modelling approach focuses on covariates which are readily available to forest managers. A two-step nonlinear mixed model was fitted to predict germination occurrence and abundance in P. pinea under varying climatic, environmental and stand conditions, based on a germination data set covering a 5-year period. The results obtained reveal that the process is primarily driven by climate variables. Favourable conditions for germination commonly occur in fall although the optimum window is often narrow and may not occur at all in some years. At spatial level, it would appear that germination is facilitated by high stand densities, suggesting that current felling intensity should be reduced. In accordance with other studies on P. pinea dispersal, it seems that denser stands during the regeneration period will reduce the present dependence on direct seeding.     

基于GM(1,1)的天然次生林空间结构预测

【目的】系统地分析已有天然次生林林分空间结构数据,通过灰色模型预测天然次生林林分空间结构各指标未来的发展趋势,对天然林经营具有十分重要的意义。【方法】以湖南大围山自然保护区典型次生林为研究对象,依据结构化森林经营理论,选取了混交度、竞争指数、角尺度、空间密度指数、开阔比数、大小比数、林分综合均质性指数作为天然次生林林分空间结构合理性评价与预测的量化指标,构建了基于GM(1,1)的天然次生林林分空间结构灰色预测模型。模型将2008年林分空间结构各指标的平均值作为初始值,并在研究区设置了面积为20 m×20 m的5个研究样地(M1,M2,M3,M4,M5),利用保护区2008-2018年典型样地林分空间信息,预测了研究区调研样地2019年-2021年林分空间结构各指标未来的变化趋势。【结果】利用精度检验机制对该模型的精确度进行了有效性检验,检验结果表明,所有指标预测合格概率P合=71.43%,良好的概率P良好=22.86%,优的概率P优=5.71%,表明该预测模型符合精度检验要求。【结论】样地未来3 a各指标整体变化尺度不大,林分空间结构基本稳定。从各指标在2008-2018年时空上的变化规律来看,各样地林分平均竞争指数、平均大小比数及平均空间密度指数是影响林分均质性指数的关键指标。     

A density management diagram for Norway spruce in the temperate European montane region

Norway spruce is one of the most important conifer tree species in Europe, paramount for timber provision, habitat, recreation, and protection of mountain roads and settlements from natural hazards. Although natural Norway spruce forests exhibit diverse structures, even-aged stands can arise after disturbance or as the result of common silvicultural practice, including off-site afforestation. Many even-aged Norway spruce forests face issues such as senescence, insufficient regeneration, mechanical stability, sensitivity to biotic disturbances, and restoration. We propose the use of Density Management Diagrams (DMD), stand-scale graphical models designed to project growth and yield of even-aged forests, as a heuristic tool for assessing the structure and development of even-aged Norway spruce stands. DMDs are predicated on basic tree allometry and the assumption that self-thinning occurs predictably in forest stands. We designed a DMD for Norway spruce in temperate Europe based on wide-ranging forest inventory data. Quantitative relationships between tree- and stand-level variables that describe resistance to selected natural disturbances were superimposed on the DMD. These susceptibility zones were used to demonstrate assessment and possible management actions related to, for example, windfirmness and effectiveness of the protective function against rockfall or avalanches. The Norway spruce DMD provides forest managers and silviculturists a simple, easy-to-use, tool for evaluating stand dynamics and scheduling needed density management actions.     

A model describing natural regeneration recruitment of Norway spruce (Picea abies (L.) Karst.) in Austria

In the absence of reliable and representative data on the frequency of seed years, seed amounts, germination and survival of seeds, the data of the observation period 1992–1996 of the permanent national forest inventory of Austria are used to develop a model describing the probability for the occurrence, density and height of Norway spruce (Picea abies (L.) Karst) natural regeneration in the forests of Austria. The forest inventory provided data on natural regeneration (saplings between 10 and 130 cm height) on clusters within a 3.89 km square sampling grid. A logistic equation is used to predict the probability for natural regeneration occurrence. Input parameters for this equation are the variables that describe the site, with slope and azimuth as continuous variables, and growth districts and vegetation types as discrete variables; the crown competition factor describes density and the quadratic mean diameter describes the stage of development of the stand. The same equation type is used to predict the probability for the occurrence of Norway spruce, conditional on the occurrence of some regeneration. An additional variable in this species specific model is a dummy variable which is set to 1 if Norway spruce occurs in the overstory and otherwise zero. Additional site variables entering this model are elevation and vegetation types characterizing soil fertility and moisture. Because the density and height of spruce regeneration depends on the stand's susceptibility to browsing, the probability for browsing is also modelled as a logistic equation depending on elevation, vegetation type, and stand density. Finally the probability distribution for height and density of the regeneration is described by two bivariate Weibull-distributions, each one describing browsed and unbrowsed Norway spruce regeneration respectively.     

Spatial prediction of forest stand variables

This study aims at the development of a model to predict forest stand variables in management units (stands) from sample plot inventory data. For this purpose we apply a non-parametric most similar neighbour (MSN) approach. The study area is the municipal forest of Waldkirch, 13 km north-east of Freiburg, Germany, which comprises 328 forest stands and 834 sample plots. Low-resolution laser scanning data, classification variables as well rough estimations from the forest management planning serve as auxiliary variables. In order to avoid common problems of k-NN-approaches caused by asymmetry at the boundaries of the regression spaces and distorted distributions, forest stands are tessellated into subunits with an area approximately equivalent to an inventory sample plot. For each subunit only the one nearest neighbour is consulted. Predictions for target variables in stands are obtained by averaging the predictions for all subunits. After formulating a random parameter model with variance components, we calibrate the prior predictions by means of sample plot data within the forest stands via BLUPs (best linear unbiased predictors). Based on bootstrap simulations, prediction errors for most management units finally prove to be smaller than the design-based sampling error of the mean. The calibration approach shows superiority compared with pure non-parametric MSN predictions.     

How does silviculture affect storm damage in forests of south-western Germany? Results from empirical modeling based on long-term observations

Storms represent the most important disturbance factor in forests of Central Europe. Using data from long-term growth and yield experiments in Baden-Wuerttemberg (south-western Germany), which permit separation of storm damage from other causes of mortality for individual trees, we investigated the influence of soil, site, forest stand, and tree parameters on storm damage, especially focusing on the influence of silvicultural interventions. For this purpose, a four-step modeling approach was applied in order to extract the main risk factors for (1) the general stand-level occurrence of storm damage, (2) the occurrence of total stand damage, and (3) partial storm damage within stands. The estimated stand-level probability of storm damage obtained in step 3 was then offset in order to describe the damage potential for the individual trees within each partially damaged stand (4). Generalized linear mixed models were applied. Our results indicate that tree species and stand height are the most important storm risk factors, also for characterizing the long-term storm risk. Additionally, data on past timber removals and selective thinnings appear more important for explaining storm damage predisposition than for example stand density, soil and site conditions or topographic variables. When quantified with a weighting method (summarizing the relative weight of single predictors or groups of predictors), removals could explain up to 20% of storm risk. The stepwise modeling approach proved an important methodological feature of the analysis, since it enabled consideration of the large number of observations without damage (“zero inflation”) in a statistically correct way. These results form a reliable basis for quantifying forest management’s direct impact on the risk of storm damage.     

Development of oaks (Quercus petraea (Matt.) Liebl.) emerged from bird-dispersed seeds under old-growth pine (Pinus silvestris L.) stands

In East Germany, there are a lot of areas covered by old pine stands. They are growing on soils, on which under natural conditions without anthropogenic impact mixed stands of oak (Quercus petraea (Matt.) Liebl.) and pine (Pinus silvestris L.), would form a natural forest. An important objective of ecological silviculture in these areas is to convert the pure pine stands into mixed oak–pine stands by using natural regeneration methods. A highly appreciated assistant in this connection is the European jay (Garrulus glandarius L.). The remarkable results of its work have been found by analyzing the natural regeneration in a few old pine stands in the forest district of Weißwasser in Saxony. Although mother trees of oak are very scarce there, natural regeneration of oak was found everywhere in the research area. The oak regeneration, undoubtedly created by the jay, amounted to at least 2000 oaks/ha. They were mainly distributed at random, while pine regeneration was aggregated in places where gaps in the pine canopy occurred. Oak regeneration was established much earlier than pine regeneration. Thus, oaks exceeded pines in age, height and diameter. The oaks were also superior to pines concerning height increment for the last three years. Thus, there is a good chance for oak to defend its prevailing role in the regeneration in the future. Probably, the next forest generation will be composed of oak and pine trees. The analysis of the quality of the oak regeneration shows that there is no substantial difference to artificially sown oak stands. This indicates that the European jay creates oak stands sufficiently both in number and quality.     

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

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

Spatial structure of genetic variation within populations of forest trees

The spatial pattern and structure of genetic variation are important aspects of the population genetics of forest stands. Combined with limits to seed and pollen dispersal, spatial structure affects the level of inbreeding and the action of natural selection. The genetic constitution of stand regeneration, following different forestry practices, is also influenced by spatial structure. For example, natural regeneration with seed trees involves sampling seed trees from a stand that may be genetically nonhomogeneous. This paper reviews theoretical and empirical results on spatial patterns of genetic variation, produced under limited gene flow and selection, in terms of recently developed spatial statistics (e.g., spatial autocorrelation). Genetic correlations in samples from spatially structured populations are also described, as well as how spatial samples can be used to characterize the structure of genetic variation, and how inferences can be made about (spatially distributed) components of fitness and yield.     

清林对云冷杉针阔混交林天然更新的影响初探

对吉林省汪清林业局金沟岭林场云冷杉针阔混交林择伐作业林分的归堆法、归带法和散铺法3种清林方式,采用标准地和路线调查法进行了天然更新调查。结果表明这3种清林方式对云冷杉针阔混交林的天然更新影响较小;散铺法对该林型林下植被的生长有影响。堆放时间超过10a的枝堆上出现了一定数量的云冷杉幼苗,这说明堆枝可以为云冷杉天然更新的发生提供更多必要条件。     

Relationships between the structural complexity and lichen community in coniferous forests of southwestern Nova Scotia

The relationships between the structural complexity of coniferous forests and the epiphytic lichen communities that inhabit them were examined in 51 conifer-dominated stands in southwestern Nova Scotia. One hundred and fifteen lichen species were studied in stands in the age range of 50–300 years. Environmental variables shaping the structural complexity of each forest stand were measured and their relationship with lichen species were assessed using a canonical correspondence analysis (CCA). The CCA revealed that the considerable variation in lichen community composition can be explained by several environmental variables associated with forest structure. The stand orientation on the first axis of the CCA found the most important variables for lichen richness to be stand age, tree stem density and snag stem density. The stand orientation on the second axis is strongly correlated with deciduous stem density and abundance including specific deciduous tree species such as Acer rubrum abundance. The analysis indicates that the greater the structural complexity in the forest, and thus the more microhabitats available, the greater the lichen species richness. These results should provide forest managers with a better understanding of the environmental variables that influence lichen diversity, and contribute to the development of more sustainable forest management strategies.     

系统收获表研究的概念和功能(英文)

传统收获表是指某一树种在特定地区和管理体系下，以立地条件和年龄为变量描述林分平均生长过程的数表．林分密度管理图是根据林分密度效果编制的可预测林分平均收获的图．通过与传统收获表和林分密度管理图的概观比较和探讨，该文系统地阐述了系统收获表的概念和功能．系统收获表的基本特点是为预测现实林分在不同立地条件和管理体系下生长过程的计算机程序．因此，系统收获表能描述在多维变量（如年龄、立地条件、林分密度、胸径和树高）条件下的现实林分和单木的无数生长过程     

Successional process of Picea crassifolia forest after logging disturbance in semiarid mountains: A case study in the Qilian Mountains,northwestern China

Selective logging is the most widely employed method of commercial timber production in Asia, and its impact on forest structure, composition, and regeneration dynamics is considerable. However, the successional processes in forest communities after logging in semiarid mountains are poorly understood. To provide more information on these processes, we used data from tree rings, direct and indirect age determinations, and field measurements of stand structure to reconstruct the historical disturbance regime, stand development patterns, and successional processes in a natural Picea crassifolia forest community in the Qilian Mountains of northwestern China. The results showed that the density of P. crassifolia forest increased significantly after logging. The densities of second growth forests 30 and 70 years after logging disturbance had increased to 2874% and 294% of primary forest's density, respectively. Logging disturbance did not alter tree species composition of logged stands. However, the diversity of understory species changed significantly among the successional phases. Logging disturbance decreased the spatial heterogeneity of second growth forest. The spatial distributions of recruitment were affected by the location of the remaining trees. There was less recruitment near the remaining trees than near forest that had been cut. In addition, logging disturbance also induced a growth release for the trees on the sites sampled. Our results imply that the succession and regeneration of P. crassifolia forest may be improved if the remaining trees could be retained relative uniform distribution pattern, thinning or selective logging could be performed to height density, exotic shrubs could be removed or the shrubs cover could be reduced during the earlier successional stages.