Tree mortality in a natural mixed forest affected by stand fragmentation and by a strong typhoon in northern Japan

Windthrow is recognized as an extremely significant disturbance in many forests. Its effects are increased by stand fragmentation, which exposes the stand to strong winds. In this study, we investigated the change in tree mortality in fragmented stands with the distance from the stand edge, in a conifer–broadleaved mixed forest in northern Japan. We set out experimental plots having three stand sizes (400, 1,600, 6,400 m²) and examined the stand dynamics over 10 years. Tree mortality tended to be higher in smaller stands, although an effect of stand size was found only in the first 5 years of the study. Distance-dependent individual mortality was obvious in Abies sachalinensis, the most vulnerable major tree species, again in the first 5 years, suggesting that wind-risk management should emphasize the area within ca. 20 m from the stand edge. No distant-dependent effect was found, however, in the latter 5 years, in which there was a strong typhoon; tree deaths occurred throughout the stand (irrespective of distance within) as a result of this event. We conclude that the severity of wind can cause the features of wind-induced damage to differ; stand edge effects peculiar to a small forest are unlikely to occur with particularly strong winds, and the effect of fragmentation might therefore be clear only in weaker disturbances.     

Inventory and analysis of tree injuries in a rockfall-damaged forest stand

Rockfall is a major threat to settlements and transportation routes in many places. Consequently, the protective function of mountain forests has recently gained particular interest. However, much is still unknown about the ideal properties of protective forest stands. Therefore the present paper discusses a method for the inventory and analysis of tree injuries in a rockfall-damaged forest stand. With this method, the interrelation between stand geometry and rockfall injuries in a subalpine Polygalo chamaebuxi-Piceetum was examined. The study site of 0.3 Ha is located in the transit zone of frequently passing, small rockfall fragments (~10 cm in diameter) causing healable tree injuries. Tree and injury parameters were recorded and analysed as to injury number, height and size. The spatial distribution of the 157 trees (diameter at breast height dbh>5 cm) in the stand as well as of the 1,704 identified rockfall injuries showed a very uneven pattern. As expected, number, height and size of the injuries generally declined with increasing distance from the cliff as well as due to higher stem densities. In contrast, results indicated that the dbh of trees has no significant influence on the number of injuries per tree. However, this study showed a clear interrelation between tree and injury distribution: in general, large trees close to the cliff and smaller trees with a high density further down the slope seem to be favourable for good protection. At least an uneven-aged, multilayered stand should be sustained. Overall, the combined analysis of stand geometry and injury parameters provides information on the spatial distribution of rockfall and on the influence of tree arrangements.     

Twenty-three years of stand dynamics in an old-growth Chamaecyparis forest in central Japan

Structures and dynamics of old-growth coniferous stands are affected by several types of disturbances including typhoons. We report the forest dynamics of four old-growth Chamaecyparis stands in central Japan that differ in the disturbance history of typhoons over a period of 23 years. The stem number, basal area and mortality were examined. In a predominant stand of C. obtusa (Sieb. et Zucc.) Endl., 24 % of the C. obtusa canopy trees died, mainly as a result of the severe damage of a strong typhoon that caused a single tree-fall gap and the following gap enlargements. In this stand, the total basal area decreased to 76.5 % of the initial value, although the mortality declined in recent years. In contrast, the other three stands decreased only slightly in the stem numbers (0.0–5.6 %) and increased in the basal areas of C. obtusa canopy trees. It is confirmed that the stand-level ingrowths of 300-year-old C. obtusa canopy trees could contribute to the increase in the stock of each stand. Our results support an idea that the dynamics of old-growth Chamaecyparis forests were greatly affected by typhoons. The stand structures will be gradually changed (with the processes of gap dynamics) and C. obtusa will continue to be dominant, potentially over hundreds of years.     

Variation in stand mortality related to successional composition

Forest ecologists have long recognized that there are large differences in the vital rates of early- and late-successional species: early-successional species typically grow faster than late-successional species, while late-successional species typically have lower rates of mortality than early-successional species. Numerous studies have shown that the differences in mortality are particularly evident when comparing per-capita rates in stands of early- and late-successional species. However, fewer studies have examined whether such differences in per-capita mortality are manifest as a difference in biomass turnover: do stands comprised late-successional species have lower rates of basal area mortality (mortality per unit basal area) than stands comprised of early-successional species? In this paper, the relationship between stand mortality and successional composition is examined using forest inventory data from the state of Michigan (USA). Mortality was quantified as the annual percentage of basal area lost to mortality, and successional composition was quantified using a continuous variable that reflects the successional status of each of the component tree species. Analysis by multivariate regression revealed that the difference in mortality between early- and late-successional stands is as great as 50%, and that the significance of this result is robust to collinearity between stand composition, stand age, and stand structure. This result suggests that successional composition could be used to better forecast changes in timber supply, habitat supply, and ecosystem function.     

Early response of stand structure and species diversity to strip-clearcut in a subtropical evergreen broad-leaved forest in Okinawa Island, Japan

Natural regeneration by strip-clearcut has rarely been used for restocking of evergreen broad-leaved forests in the East Asia. To contribute to a better understanding of the likely reasons for either the successful recovery of these forests or failure, the early response of stand structure and tree species diversity to strip-clearcut were studied in Okinawa Island, southwest Japan. The 4,000 m² study area was divided into 10 adjacent sections of equal area with a 10 m × 10 m sampling plot in the center, of which five sections were strip-clearcut while the remaining five sections uncut. Eight-year secondary succession following strip-clearcut showed that the density of woody stems, tree species diversity increased significantly not only in the regenerating stands but also in the residual stands compared to the primary stands. The primary dominant target species, Castanopsis sieboldii, continued to dominate both the regenerating and residual stands although colonizing species invaded abundantly in the stands, occupying approximately one-third of total tree species in each treatment. The community similarity analyses showed that the species composition were similar between the primary stands and the regenerating stands or residual stands reflected the high similarity amongst the primary stands and the regenerating stands or the residual stands. The similar stand structure, high tree species diversity and greater community similarity after strip-clearcut provided no evidence of forest degeneration, suggesting that the regenerating stand might gradually develop into stand similar to that prior to strip-clearcut.     

Spatial organisation of a bimodal forest stand

Many populations have a bimodal size distribution, even when composed of a single cohort. In developing forest stands, this pattern is usually attributed to asymmetric competition at canopy closure among trees which have access to the upper canopy and those which have failed to reach it. Nevertheless, alternative explanations for bimodality exist, and in sessile organisms spatial pattern analysis can be used to compare their predictions. A 0.25-ha plot was created in a maturing stand of Asiatic white birch (Betula platyphylla Sukacz.) in Central Kamchatka. All stems >1 cm diameter at breast height (DBH) were fully mapped. Mark correlation analysis revealed size compensation among stems up to 3.5 m apart, providing evidence that competition affected the distribution of stem sizes. The spatial pattern of trees was analysed using the pair correlation function g(r). Large trees (>20 cm DBH) had a dispersed distribution to which a Strauss soft-core Gibbs process model was fitted. This suggested that large trees interacted at scales up to 4.16 m. Small trees (1–20 cm DBH) were distributed randomly, but a cross-pair correlation analysis revealed a greater likelihood of occurrence beginning at 4.3 m from large trees, closely matching the modelled interaction distance. These results are consistent with the hypothesis that asymmetric competition is structuring this bimodal cohort of trees: large stems tend towards a dispersed pattern, exerting competitive effects at scales up to approximately 4 m, whereas smaller stems are more commonly found in the interstices within the pattern of large trees.     

Study on dynamics of stem form of Korean pine trees in natural forest stand

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Spatial variability in stand structure and density-dependent mortality in newly established post-fire stands of a California closed-cone pine forest

Fire is an important process in California closed-cone pine forests; however spatial variability in post-fire stand dynamics of these forests is poorly understood. The 1995 Vision Fire in Point Reyes National Seashore burned over 5000 ha, initiating vigorous Pinus muricata (bishop pine) regeneration in areas that were forested prior to the fire but also serving as a catalyst for forest expansion into other locales. We examined the post-fire stand structure of P. muricata forest 14 years after fire in newly established stands where the forest has expanded across the burn landscape to determine the important factors driving variability in density, basal area, tree size, and mortality. Additionally, we estimated the self-thinning line at this point in stand development and compared the size-density relationship in this forest to the theorized (−1.605) log-log slope of Reineke’s Rule, which relates maximum stand density to average tree size. Following the fire, post-fire P. muricata density in the expanded forest ranged from 500 to 8900 live stems ha⁻¹ (median density = 1800 ha⁻¹). Post-fire tree density and basal area declined with increasing distance to individual pre-fire trees, but showed little variation with other environmental covariates. Self-thinning (density-dependent mortality) was observed in nearly all stands with post-fire density >1800 stems ha⁻¹, and post-fire P. muricata stands conformed to the size-density relationship predicted by Reineke’s Rule. This study demonstrates broad spatial variability in forest development following stand-replacing fires in California closed-cone pine forests, and highlights the importance of isolated pre-fire trees as drivers of stand establishment and development in serotinous conifers.     

Diachronic analysis of individual-tree mortality in a Norway spruce stand in the eastern Italian Alps

? Understanding tree mortality processes across time requires long term studies. Spatiotemporal patterns of mortality in a 200 years-old mono-layered Norway spruce stand were evaluated to determine what factors affected individual-tree mortality.

? We performed an analysis on two surveys (1993 and 2005) in a 1-ha permanent plot in the Paneveggio forest (Eastern Italian Alps). Tree diameter and age distribution between surveys were compared. We examined spatial patterns of living and dead trees before 1993, in 1993 and in 2005 using univariate and bivariate Ripley’s K(d) function, and a kernel estimator of local crowding. A logistic model was used to assess the effects of diameter, age, recent growth and competitive pressure on tree mortality.

? Spatial pattern analysis indicated mortality was associated to tree neighbourhood (neighbour effect at 2–5 m). An increment of regularization of tree spatial pattern occurred due to density-dependent mortality. Logistic regression showed tree diameter and recent growth were determinant on mortality risk during the monitoring period.

? Even if the stand is relatively aged, mortality dynamics are those typical of stem exclusion stage. Mortality was related to competitive dynamics, and small suppressed trees with slow growth rate had higher probability to die.

     

Belowground competition in a broad-leaved temperate mixed forest: pattern analysis and experiments in a four-species stand

We investigated fine root biomass and distribution patterns in a species-rich temperate Carpinus–Quercus–Fagus–Tilia forest and searched for experimental evidence of symmetry or asymmetry in belowground competition. We conducted extensive root coring and applied the recently introduced in situ-root growth chamber technique for quantifying fine root growth under experimentally altered intra- and interspecific root neighbourhoods in the intact stand. In 75% of all soil cores, fine roots of more than two tree species were present indicating a broad overlap of the root systems of neighbouring trees. Quercus trees had more than ten times less fine root biomass in relation to aboveground biomass or productivity (stem growth) and a much higher leaf area index/root area index ratio than Carpinus, Fagus and Tilia trees. The root growth chamber experiments indicated a high belowground competitive ability of Fagus in interspecific interactions, but a low one of Quercus. We conclude that (1) interspecific root competition is ubiquitous in this mixed stand, (2) root competition between trees can be clearly asymmetric, and (3) tree species may be ranked according to their belowground competitive ability. Fagus was found to be the most successful species in belowground competition which matches with its superiority in aboveground competition in this forest community.     

Changes in aboveground and belowground properties during secondary natural succession of a cool-temperate forest in Japan

Forest development in temperate regions is considered to be a global carbon sink. Many studies have examined forest development after harvesting or fire from aboveground (e.g., biomass) or belowground (e.g., soil nutrient) perspectives. However, few studies have explored forest development from both perspectives simultaneously in cool-temperate forests in Japan. In this study, we examined changes over 105 years in both aboveground and belowground components during secondary natural succession. The aboveground biomass increased for 50 years and reached a plateau in a 105-year-old stand. The N mineralization rate increased during succession for 50 years, but showed a decline in the 105-year-old stand due to the decrease in the nitrification rate in late succession. The percent nitrification (i.e., relative contribution of nitrification to N mineralization) decreased significantly with increasing forest stand age. The N mineralization rates had significant relationships with N concentrations of the dominant tree foliage and litter fall and with the amount of litter fall N. Meanwhile, other belowground properties (i.e., soil pH, phenol concentration, soil microbial respiration, and litter mass loss) did not show any significant relationship with forest stand age. This may be because the soil at the study sites was heterogeneous and consisted of Cambisols and Andosols, the latter of which originally has high organic matter content, and thus may have buffered the effect of the aboveground development. These results indicate that belowground N dynamics are more closely associated with aboveground development than other belowground properties in these forests.     

Response of a rising 2-year-old natural Piedmont upland stand to fertilization

Hardwood stands in the southern U.S. are often regenerated naturally following clearcutting, with little or no silvicultural intervention in the early stages of stand development. Fertilizer was applied to a young naturally regenerating stand in order to evaluate the effectiveness of nutrient addition as a silvicultural tool in recently clearcut stands and to better understand the ecological relationships between site fertility and stand development. The study was installed on a rising 2-year-old naturally regenerated mixed pine-hardwood stand in the Hill Demonstration Forest in the Piedmont region (Durham County) of North Carolina. Fertilizer treatments were control, N, N + P, and N + P + K (at 200 kg N per ha, 50 kg P per ha and 100 kg K per ha). On a whole stand basis, increased growth rates were observed for trees on N + P and N + P + K plots. Fertilizer treatments did not affect total stand density, but the density of loblolly pine significantly decreased in N + P plots compared to the Control. The density of stems of stump origin also increased in plots receiving N + P. Density of sweetgum (Liquidambar styraciflua) significantly increased and hickory (Carya spp.) density decreased with the application of N + P. Dominant hardwood species with the exception of yellow-poplar (Liriodendron tulipifera) responded with an increase in height to only N + P fertilizer treatments. Yellow-poplar increased in groundline diameter, height and mean tree volume index with the application of N + P. Loblolly pine (Pinus taeda) responded to N + P + K fertilizer treatment with an increase in height, diameter and volume index over the Control. There was no growth response among treatments for the 16 largest trees per species per plot, with the exception of red oak (Quercus rubra) which responded with an increase in height in N + P plots. Projected growth responses were modeled on a whole stand level and indicate that the use of N + P fertilizer may be an effective silvicultural instrument to increase growth, accelerate stand development, and shorten rotation length.     

Improving the simulation of stand structure in a forest gap model

There is currently great interest in improving the applicability of forest gap models to changing environmental conditions, in order to facilitate the assessment of possible impacts of climatic change on forest ecosystems. Moreover, for the development of mitigation strategies, it is necessary to include forest management options in the models. Both the simulation of transient effects of climatic change and of forest management regimens require a realistic representation of stand structure in gap models, since tree species respond to variations in stand density in characteristic ways, depending on their ecological strategies.In this study, we compared the effect of five different height growth functions that are sensitive to stand density on simulated stand structure of the FORSKA forest gap model. We used long term observation data from a beech thinning trial at Fabrikschleichach, Bavaria, to test the alternative functions. First, we compared simulation results of the original FORSKA model with measured stand development from 1870 to 1990. Whereas simulated stand level variables (e.g. biomass, mean diameter and height) showed good correspondence with observations, individual tree dimensions and simulated stand structure were quite unrealistic. After calibrating parameters of the height growth functions with data from a lightly thinned plot at Fabrikschleichach, we ran the model with data from a heavily thinned plot for validation. All five functions considerably improved the simulation of height/diameter relationships and stand structure. However, there were distinct differences between functions. The best correspondence with measurements was shown by a function which uses the relative radiation intensity in the centre of a tree crown as an indicator of the competition status of the tree. This function is rather simple and needs only two growth parameters, which can be derived for different functional types of species, according to their shade tolerance.With the new, flexible height growth function it should be possible to extend the applicability of gap models to more realistic simulation experiments including forest management and natural disturbance. To our knowledge, this was the first attempt to employ long term forest observation data for the calibration and validation of a forest gap model. The results suggest that such data could be very useful in model testing and improvement.     

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。     

13C and 15N isotopic fractionation in trees, soils and fungi in a natural forest stand and a Norway spruce plantation

¹⁵N and ¹³C natural abundances of foliage, branches, trunks, litter, soil, fungal sporophores, mycorrhizas and mycelium were determined in two forest stands, a natural forest and a Norway spruce plantation, to obtain some insights into the role of the functional diversity of saprotrophic and ectomycorrhizal fungi in carbon and nitrogen cycles. Almost all saprotrophic fungi sporophores were enriched in ¹³C relative to their substrate. In contrast, they exhibited no or very little shift of δ¹⁵N. Judging from the amount of C discrimination, ectomycorrhizal fungi seem to acquire carbon from their host or from dead organic matter. Some ectomycorrhizal species seem able to acquire nitrogen from dead organic matter and could be able to transfer it to their host without nitrogen fractionation, while others supply their host with ¹⁵N-depleted nitrogen. Moreover ectomycorrhizal species displayed a significant N fractionation during sporophore differentiation, while saprotrophic fungi did not.     

伞伐更新法对赫卡尼亚(Hyrcanian)森林系统中森林再生和林分结构的影响

研究伞伐更新法对赫卡尼亚森林系统中森林再生和林分结构的影响。在伊朗北部优势种为山毛榉（Fagus orientalisLipsky）的硬木林,采取伞伐更新法和非伞伐更新法方法设计实验区。结果表明,在山毛榉（F orientalis Lipsky）林中采用伞伐更新法处理,明显影响林下草本植物种的频度和密度。伞伐更新法处理后,林堇菜（Viola silvestris Lam.）、车叶草（Asperula odorata L.）、苔草（Carex spp.）和悬钩子（Rubushyrcanus Juz）的种频度明显增加。在对照区,树种的胸高直径（57.50±2.15cm）大于被处理区（50.67±1.88cm）的树种的胸高直径（50.67±1.88 cm）,但对照区和处理区的树种的高度值相似。在1995-2005年间,波斯铁木（Parrotia persica）苗木数量增加到13.2%,而山毛榉（F orientalis）和欧洲鹅耳枥（Carpinus betulus）的苗木数量却明显减少。总之,应该利用其它育林方法,如,带状择伐作业,而不是伞伐更新法培育赫卡尼亚森林中山毛榉林。     

Wood production in a natural stand of Acacia karroo in Zimbabwe

Trees are important components of agricultural systems in the dry tropics of Africa. The assumption that exotic species are more productive than the indigenous woodland in these situations is seldom supported by quantitative data. In this study a new technique of recognising annual growth rings from the bands of marginal parenchyma has been used to model growth in a natural population of Acacia karroo Hayne, one of the most widespread and useful trees in the savanna woodlands of southern Africa. The study showed both basal area and sectional area at ankle height to be excellent predictors of total tree volume. Stands may yield from about 1 to 4 m³ ha⁻¹ year⁻¹ dependent upon spacing and site; individual trees produced up to 1 m³ of wood in 26 years. Maximum current annual increment was reached about 12 years after the trees approached breast height, thereafter declining until they became moribund about 8 years later. The economic rotation for wood production was close to the lifespan of the tree.     

广东四会贞山森林旅游区林分改造造林效果初探

根据广东省四会市大南山林场贞山林区生态林的现状及所处的特殊地理位置状况,按照建设森林公园、提高贞山风景区生态功能等级和景观质量的目标,规划设计了森林旅游区首期林分改造的树种选择及配置,初步分析了林分改造各树种的生长情况,展望了改造后的生态、景观前景,对今后林分改造提出建议。     

Effects of topography and management history on natural forest recovery in abandoned forest after clear-cutting in Miyazaki, Japan

We investigated factors limiting the recovery of natural forest in former large-scale conifer plantations abandoned after clear-cutting in southwestern Japan. We analyzed forest recovery status (“recovered” sites covered by evergreen broad-leaved trees, and “unrecovered” sites covered by pioneer community or nonvegetated sites) using aerial photographs and field survey. We applied logistic regression analyses to evaluate the effects of topography, construction of harvesting roads, distance from remnant forest, stand condition before clear-cutting, and prior land-use history on forest recovery. Human factors, i.e., land use and clear-cutting age, were found to affect to forest recovery more than environmental factors such as topography. Harvesting roads had the strongest negative impact on forest recovery. Forest recovery after clear-cutting of young sugi plantations also took longer than after clear-cutting of old sugi plantations or evergreen broad-leaved forests. Furthermore, areas formerly utilized as meadows recovered less successfully than those that had been managed as coppices. The influences of these factors were thought to be promoted by the advance reproduction as the regeneration sources for forest recovery. The influence of stand age before logging suggested an effect of thinning, which might alter the abundance of advanced reproduction in the understory. However, distance from remnant forest appeared to be less important. An influence of topography was also detected, but this could be partly explained by the existence of advance reproduction in the understory in certain topographic positions. Thus, our analysis suggests that regeneration sources originating from advanced reproduction in plantations play a significant role for the recovery of natural forest after clear-cutting.