基于竞争指数的杉木林分生长可视化模拟研究

以我国南方速生丰产林——杉木纯林为研究对象,采用改进过的Hegyi的简单竞争指数模型对杉木纯林间的竞争强度进行定量分析,并建立单木生长模型,再通过径阶模型将其应用于全林分,将单木生长模型研究扩展到全林分生长模型研究。同时,通过可视化模拟技术,为更加逼真的恢复和重建林分空间结构提供理论依据。     

Modelling the edge effect in even-aged Monterey pine (Pinus radiata D. Don) stands

Six edge effect models are presented. Modelling results for the edge effect on the DBH, total height, crown height, and basal area in even-aged stands of Monterey pine (Pinus radiata D. Don) are provided. Free edges and edges under external competition pressure were analysed. Model fitting was carried out in two phases. In the first phase, independent fittings for each border type by sampled cardinal point in each stand were carried out to detect differences in edge effect intensity and depth according to edge aspect. In the second fitting phase, the best models selected by edge aspect in the first phase were restricted to estimate the same value for each variable in the stand interior, independently of the border where it was analysed.

The edge effect intensity and depth varied depending on the analysed variable and edge type. Modelling of the edge effect on DBH and total height on free edges of old stands requires a model with independent parameters for each border aspect; the models can be restricted to estimate the average value of each variable in stand interior. A model with independent parameters for each border aspect should be also used for modelling the effect on total height, crown height and basal area for the edges of young stands under external competition pressure. The modelling of basal area and crown height on free edges of old stands, DBH on edges of young stands under external competition pressure, and all variables mentioned on free edges of young stands can be carried out through an average fitting. DBH and basal area are directly affected by stand density; thus, modelling of edge effect on these and others variables which are affected by stand density can improve with the inclusion of a competition index or a point density index.     

乳源木莲杉木混交林生长及其竞争关系分析

阔叶林尤其是常绿阔叶林是亚热带地区的地带性植被,它的存在对维护我国广大南方地域的生态平衡、满足经济建设和人民生活需要起着极其重要的作用。但近年来,阔叶林的人为破坏及阔叶树种人工培育不受重视,加上发展针叶人工林,致使针阔比例严重失调,产生一系列生态问题,很多优良乡土阔叶树种随着天然林的肆意砍伐利用而逐步消失(盛炜彤,1993;俞新妥,1997;陈存及等,2000)。为了扭转这种局面,福建省近年来开始重视阔叶树种的造林,大力挖掘优良乡土树种和寻找新树种,特别是珍稀速生阔叶树种。但是很多优良乡土树种连采种育苗都极为困难,乳源木莲…     

Individual tree growth response to variable-density thinning in coastal Pacific Northwest forests

We examined 5-year basal area growth of nearly 2600 trees in stem-mapped plots at five locations differing in site characteristics, species composition, and management history on the Olympic Peninsula in Western Washington, USA. Our objectives were to determine if internal edges, the boundaries within the stand between components of the variable-density thinning, influenced individual tree growth, and whether incorporation of individual tree local competition indices in growth prediction models could account for treatment and edge effects. Treatment significantly affected tree growth at all sites, with trees in the thinned matrix displaying on average over 25% greater basal area growth than trees in unthinned patches. Proximity to canopy gaps created as part of the variable-density thinning increased basal area growth of trees in the thinned matrix by nearly 11%. In addition, growth of trees close to skid trails was 11% greater than trees located away from the trails. Past thinning history, and its effect on initial stocking rate, appeared to affect the magnitude of the edge effects. Blocks that had received earlier commercial thinnings, and thus had lower stocking at the onset of the study, displayed lower growth responses than previously unthinned blocks. Including local competition indices in the models generally reduced growth prediction error; however, the indices examined did not fully account for treatment or edge effects. Our results suggest that not accounting for internal edges in spatially complex stands could result in errors in projected growth of trees, although these edge effects are highly variable. Failure to account for the effects of internal edges could affect not just estimates of future stand yield, but also projections of future stand structure.     

甘肃连城国家级自然保护区青海云杉种内种间竞争的研究

采用Hegyi提出的单木竞争指数模型对甘肃连城国家级自然保护区的青海云杉种内、种间的竞争指数进行定量分析。结果表明:青海云杉种内竞争指数随着林木径级的增大而逐渐减小,种内竞争远远大于种间竞争,种内种间竞争指数的大小顺序为青海云杉>青杄>桦叶四蕊槭>山杨>红桦>柳树;竞争指数与对象木的胸径之间服从幂函数关系CI=AD~B,当青海云杉胸径达到45cm以上时,竞争指数变化很小;青海云杉的竞争指数与海拔和郁闭度呈显著正相关性。     

Growth and nut production of black walnut in relation to site, tree type and stand conditions in south-central United States

Black walnut (Juglans nigra) is a prime tree species for agroforestry practices in the United States providing highly prized wood and nuts for human consumption and wildlife. In 54 black walnut stands in south central United States, the site index (i.e., mean dominant height [DH], at age 25 years) ranged between 5.2 and 21.4 m, and was independent of stand density. There were no differences in height and stem diameter (DBH) growth rates between stands with improved varieties and native stock. Most stands were in a “free growth” stage because of either early age or wide spacing. Mean annual increments in DBH and height were positively related both for improved varieties and native stock. Understory competition had a substantial detrimental effect on DH. In a 26 year-old stand, trees growing within Kentucky blue grass (Poa pratensis) had a site index 5 m greater than trees growing within tall fescue (Festuca arundinacea). Predicted nut yields in 2002 varied between 0 and 1370 kg of hulled nuts per ha. Improved varieties had, in general, higher nut yields than predicted from a nut yield-DBH equation developed for individual trees. Nut yields were highly variable both within and among stands, and were related to DBH in native stock but not in improved varieties.     

Edge type affects leaf-level water relations and estimated transpiration of Eucalyptus arenacea

While edge effects on tree water relations are well described for closed forests, they remain under-examined in more open forest types. Similarly, there has been minimal evaluation of the effects of contrasting land uses on the water relations of open forest types in highly fragmented landscapes. We examined edge effects on the water relations and gas exchange of a dominant tree (Eucalyptus arenacea Marginson & Ladiges) in an open forest type (temperate woodland) of south-eastern Australia. Edge effects in replicate woodlands adjoined by cleared agricultural land (pasture edges) were compared with those adjoined by 7- to 9-year-old eucalypt plantation with a 25m fire break (plantation edges). Consistent with studies in closed forest types, edge effects were pronounced at pasture edges where photosynthesis, transpiration and stomatal conductance were greater for edge trees than interior trees (75m into woodlands), and were related to greater light availability and significantly higher branch water potentials at woodland edges than interiors. Nonetheless, gas exchange values were only ～50% greater for edge than interior trees, compared with ～200% previously found in closed forest types. In contrast to woodlands adjoined by pasture, gas exchange in winter was significantly lower for edge than interior trees in woodlands adjoined by plantations, consistent with shading and buffering effects of plantations on edge microclimate. Plantation edge effects were less pronounced in summer, although higher water use efficiency of edge than interior woodland trees indicated possible competition for water between plantation trees and woodland edge trees in the drier months (an effect that might have been more pronounced were there no firebreak between the two land uses). Scaling up of leaf-level water relations to stand transpiration using a Jarvis-type phenomenological model indicated similar differences between edge types. That is, transpiration was greater at pasture than plantation edges in summer months (most likely due to greater water availability at pasture edges), resulting in significantly greater estimates of annual transpiration at pasture than plantation edges (430 vs. 343lm(-2)year(-1), respectively). Our study highlights the need for landscape-level water flux models to account for edge effects on stand transpiration, particularly in highly fragmented landscapes.     

日本落叶松人工林密度调控技术研究

以日本落叶松中幼林抚育间伐样地监测数据,利用树冠竞争因子（CCF）、林分密度指数、单位断面积生长过程等指标,分析了抚育间伐对断面积和蓄积两方面的影响,结果表明：间伐林分的断面积和蓄积有相同的生长过程,即林分在一定间伐强度范围内（强度小于50％）间伐强度越大,年均生长率越大;用树冠竞争因子描述日本落叶松人工林断面积的生长过程,未间伐林分的立木度接近稳定状态,而间伐林分降低了林分的CCF值,随着林龄的增加,CCF值不断增加,逐渐向未间伐林分靠近。总的来说间伐林分在间伐初期要小于未间伐林分,但随着林龄的增大及林木竞争的加剧,间伐林分逐渐向未间伐林分靠近。     

Modelling growth behaviour in monoculture in subtropical eastern Australia of two eucalypt species that differ in shade tolerance

Growth behaviour of the relatively shade-intolerant species Eucalyptus dunnii and the rather more shade-tolerant Corymbia citriodora was considered during several growth periods over 8–14 years of age in experimental plantation monoculture in subtropical New South Wales. Larger trees dominated growth in any stand; this was consistent with the principle that inter-tree competition in such forests is for light and is asymmetric in that larger trees shade smaller trees but the reverse does not occur. Once competition was established, competition-induced mortality of C. citriodora was less than that of E. dunnii. A model relating tree diameter growth rate to tree diameter, competitive position in the stand and stand basal area was developed. The model showed that smaller, hence partially shaded, trees in stands of C. citriodora were better able to maintain their growth rates than equivalent trees in stands of E. dunnii. It is hypothesised that this reflects the greater shade tolerance of C. citriodora. This pattern of growth led to a more even distribution of tree size classes across the stand in the more shade-tolerant species.     

On the influence of windward edge structure and stand density on the flow characteristics at forest edges

Experimental investigations in an atmospheric boundary layer wind tunnel were carried out in order to study both the influence of the windward edge structure and the stand density on the flow characteristics near the canopy top of forest edges. For the edge structure investigation, two different types of forest edges were used. The taper angle was varied three times for both edge types, and all tapered edges were combined with both dense and sparse forest stands. In addition, a configuration was investigated where the airflow in the trunk space of the dense forest was blocked completely by an impermeable wall. In order to study the influence of stand density, forest stands were varied three times by removing whole rows of the originally dense model forest. The measurements show that a forest edge induces strong disturbances in the flow field at the near-edge region leading to an increased momentum transfer and to an increased wind loading on trees. The region of significant disturbances extends to a distance of about eight tree heights. Both edge density and taper angle determine the amount of volume influx through the edge, which seems to be primarily responsible for the flow field properties near the canopy top. It was found that for sparse forest stands, sloping tapered edges reduce the wind velocities nearest to the canopy. It could also be shown that the positive skewness of the near-canopy longitudinal velocity distribution is diminished with decreasing taper angle indicating that higher wind velocity fluctuations are reduced.     

福建光泽天然林中南方铁杉的种内与种间竞争

通过各种竞争指数的比较,提出采用Hegyi单木竞争指数模型研究福建光泽天然林中南方铁杉在生长发育过程中受到的种内和种间竞争强度。结果表明,南方铁杉受到的种内竞争指数为52.03,高于任一其他树种的竞争指数,但低于种间竞争指数总和(164.08)。南方铁杉受到的种间和种内竞争大小顺序为:南方铁杉>木荷>猴头杜鹃>甜槠>棕脉花楸>小叶青冈>鹿角杜鹃>厚叶红淡比>黄山松>鼠刺叶石栎>野漆>吴茱萸五加>厚叶厚皮香>柳杉>枫香>榕冬青>石灰花楸>马醉木>树参>冬青>银钟花。南方铁杉对象木受到的竞争强度随着对象木径级的增大呈明显减少趋势。当对象木胸径大于30 cm后,个体受到的竞争强度趋于稳定。竞争指数与对象木胸径的关系呈幂函数关系。     

A hypothesis about the interaction of tree dominance and stand production through stand development

The development of forests over time involves changes in rates of growth of trees and stands, and changes in the competition and dominance between trees plays a large role in the overall development of stands. A hypothesis proposes that changes in the growth of trees and stands result from regular changes in dominance and the efficiency of resource use by dominant and non-dominant trees. Dominance is low prior to canopy closure, and efficiency of resource use is high for all trees. Increasing dominance near canopy closure reduces the efficiency of resource use by non-dominant trees, lowering overall stand growth. Later in stand development, the efficiency of resource use also declines for the largest trees, reducing the level of dominance in the stand. The dominance part of this hypothesis was examined for 150 years of stand development in two mixed-species stands in the Coast Range of Oregon. A quantitative index of dominance was minimal prior to the peak in stand growth near age 25–30 years, and then increased sharply as stand productivity declined. Dominance then declined after age 100 years as the growth rate of the 300 largest trees/ha began to decline. The dominance portion of the hypothesis was supported, and further testing may be useful.     

Effects of residual trees on tree height of 18- and 19-year-old Cryptomeria japonica planted in group selection openings

This study examined the effects of residual trees on the height of 18- and 19-year-old Cryptomeria japonica planted in group selection openings of about 0.1 ha at Yufuin in Oita Prefecture in southern Japan. We first developed a general model expressing variation in the height for all of the planted trees within the openings from indices of both between-cohort competition (the effects of residual trees) and within-cohort competition (among the even-aged planted trees). The between-cohort competition index was calculated both with and without taking into account the direction that residual trees were positioned from subject trees. The two models suggested that residual trees located near the northern edge of the opening were not effective competitors for light for the planted trees within the opening. In addition, our results indicated that both between-cohort competition and within-cohort competition are important for explaining tree height in closed stands within group selection openings. Using the general model, we then estimated the potential loss of height growth for locally dominant trees within the openings that could be attributed only to the influence of residual trees. These results indicated that in addition to the effect of opening size, both the degree of crowding and the direction of residual trees from the planted trees are important factors affecting the height of individual trees planted in group selection openings.     

Models for predicting height and diameter of individual trees in young Picea abies (L) karst. stands

Models for predicting height and diameter of individual trees in young Picea abies (L.) Karst. stands were developed. Data collected in a large survey of young forest stands in Sweden (the HUGIN young stand survey) were used in the construction of the models. Models were developed both with and without competition indices included. When constructing the competition indices trees within three metres from the subject tree were regarded as competitors. Functions with competition indices included (distance dependent) will be useful in analyses of the development of stands with different stand structure, whereas functions without competition indices (distance independent) will be useful in systems for long‐term forecasts of yield.     

Spatial structure of deciduous forest stands with contrasting human influence in northwest Spain

Five contrasting deciduous forest stands were studied to characterize the spatial structural variability in human-influenced forests. These stands are representative of cultural forest types widely represented in western Europe: one plantation, two coppices, one wood-pasture forest and one high forest stand. All stems with DBH > 5 cm were measured and mapped, and stem DBH distributions, spatial structure of DBH, spatial point patterns and spatial associations were analysed. Spatial autocorrelation for DBH was calculated with Moran’s I correlograms and semivariograms. Complete spatial randomness hypothesis for spatial point patterns, and both independence and random labelling hypotheses for spatial associations were analysed using Ripley’s K function. The results showed that tree sizes were conditioned by particular former management systems, which determined unimodal symmetric, positively skewed or compound DBH distributions. Spatial structure was more complex when human influence became reduced. Coppice stands showed clumped spatial patterns and independence among size classes, as a consequence of sexual and vegetative establishment of new stems in open areas. The largest clumping intensity was observed in the wood-pasture with an intermediate disturbance frequency and low inter-tree competition. The high forest stand displayed spatial traits consistent with the gap-dynamics paradigm, such as clumping of smaller trees, random arrangement of larger trees, negative association between juveniles and adults, and high structural heterogeneity. It can be expected that after cessation of human interference, coppices and wood-pastures would evolve to a more heterogeneous structure, probably with a higher habitat and species diversity.     

Individual-based measurement and analysis of root system development: case studies for <Emphasis Type="Italic">Larix gmelinii</Emphasis> trees growing on the permafrost region in Siberia

We present results of individual-based root system measurement and analysis applied for Larix gmelinii trees growing on the continuous permafrost region of central Siberia. The data of root excavation taken from the three stands were used for the analyses; young (26 years old), mature (105 years old), and uneven-aged over-mature stand (220 years old). In this article, we highlight two topics: (1) factors affecting spatio-temporal pattern of root system development, and (2) interactions between aboveground (i.e., crown) and belowground (i.e., root) competition. For the first topic, the detailed observation of lateral roots was applied to one sample tree of the overmature stand. The tree constructed a superficial (<30 cm in depth) and rather asymmetric root system, and each lateral root expanded mainly into elevated mounds rather than depressed troughs. This indicated that spatial development of an individual root system was largely affected by microtopography (i.e., earth hummocks). For these lateral roots, elongation growth curves were reconstructed using annual-ring data, and annual growth rates and patterns were compared among them. The comparison suggested that temporal root system development is associated with differences in carbon allocation among the lateral roots. For the second topic, we examined relationships between individual crown projection area (CA) and horizontal rooting area (RA) for the sample trees of each stand. RA was almost equal to CA in the young stand, while RA was much larger (three or four times) than CA in the mature and overmature stands. Two measures of stand-level space occupation, crown area index (aboveground: CAI; sum of CAs per unit land area) and rooting area index (belowground: RAI; sum of RAs), were estimated in each stand. The estimates of RAI (1.3–1.8 m² m₋₂) exceeded unity in all stands. In contrast, CAI exceeded unity (1.3 m² m₋₂) only in the young stand, and was much smaller (<0.3 m² m₋₂) in the two older stands. These between-stand differences in RAI–CAI relationships suggest that intertree competition for both aboveground and belowground spaces occurred in the young stand, but only belowground competition still occurred in the two older stands. Based on this finding, we hypothesized that competition below the ground may become predominant as a stand ages in L. gmelinii forests. Methodological limitations of our analysis are also discussed, especially for the analysis using the two indices of space occupation (CAI, RAI).     

海南霸王岭陆均松类热带山地雨林伐后林结构与物种多样性研究

霸王岭林区是海南岛最典型的热带山地雨林分布区.在该区,现存分布面积最广、资源数量最大的是热带山地雨林的各种伐后林.以霸王岭分布最广的以陆均松(Dacridium pierrii)为主的山地雨林的伐后林为对象,研究了其组成、径级、高度结构和物种多样性特征,对比分析了伐后林与原始林以及伐后林不同类型之间的异同.结果表明:原始热带山地雨林经采伐后中小乔木和灌木的优势度增大,而大乔木树种的优势度则明显下降.阳性树种的种类、数量和相对重要性随采伐破坏的程度而增加.伐后林中矮、小树木在林分中比例随采伐破坏的程度增加,而高、大树木,特别是进入主林层树木的比例则随采伐破坏的程度明显下降.从林分测树因子的变化来看,采育择伐和径级择伐的林分要优于大强度择伐和皆伐的林分.原始林的物种丰富度大于伐后林,但其多样性指数和均匀度指数大都小于伐后林.组成上与原始林相似性较大的伐后林是各种择伐后的林分,而皆伐后的林分与原始林在组成上已有很大差别.     

Using a retrospective dynamic competition index to reconstruct forest succession

Understanding forest dynamics and stand structures is crucial for predicting forest succession. However, many forests have been altered due to century-long land-use practices, which complicates the reconstruction of past and current successional trajectories. For a better understanding of successional processes, we suggest studying the intra- and interspecific competition among single trees across time. We introduce a tree-ring based competition index to reconstruct the competitive dynamics of individual trees over time. This new retrospective dynamic competition index combines a temporal and a spatial component by calculating the yearly ratio between the basal area increments (bai) of the neighbouring trees and the subject tree. The new index is applied to mixed Scots pine (Pinus sylvestris L.) and pubescent oak (Quercus pubescens Willd.) stands in the inner-Alpine dry-valley Valais, for which a change in species composition is hypothesised. The aim is to analyse current stand structures in terms of recent changes in the competitive interactions at the single tree level and to relate these competitive dynamics to land-use change and increasing drought due to climate change. On five plots, the positions of 456 trees were recorded and increment cores were taken to derive bai data. The individual dynamic competition index curves were aggregated in clusters, which define typical patterns of competitive dynamics in both tree species. A large percentage of the trees (87% in oak, 70% in pine) were clustered into a group of trees with constant competition at a relatively low level. However, a smaller group of pines (20%) had recently faced increasing competition. In addition, stand structure analyses indicated a change towards a higher proportion of oak. This change in the competitive ability between oak and pine was found to be related to drought, in that oak had a competitive advantage in dry years. Furthermore, the high proportion of dead branches in pines with decreasing competitive abilities indicated increasing competition for light as a consequence of natural development towards a later successional stage that favours the more shade-tolerant oak. The new retrospective dynamic competition index proved to be promising in studying forest succession. The tree-ring based method allows us to identify changes in the competitive ability of single trees with a high temporal resolution and without repeated assessments.     

用二类调查样地建立落叶松单木直径生长模型

根据来自吉林省汪清林业局二类调查的１４个落叶松固定样地的３４６株复测林木数据,建立了与我关的落叶松木平方直径生长最模型。它包含了单木直径生长模型所需考虑的林木大小,立地条件和林木竞争这３个因素,并具有与年龄、地位指数无关的特点,便于实践中应用,逐步回归的结果表明,落叶松单木平方直径生长最模型需要考虑的主要因子是林木直径值、林分断面积和坡度。     

Influence of umbrella pine (<Emphasis Type="Italic">Pinus</Emphasis><Emphasis Type="Italic">pinea</Emphasis> L.) stand type and tree characteristics on cone production

Most umbrella pine (Pinus pinea L.) stands are managed as agroforestry systems, whose main production is fruit, due to the edible and highly nutritious kernels, and are frequently associated to natural or seeded pastures and grazing. The stands have low density, in order to enhance crown growth and fruit production. Nevertheless, cone production, both with regard to number and weight, varies greatly between stands, trees and years. In this study were selected three agroforestry systems, representative of umbrella pine stands whose main production is fruit, and one stand representative of the timber production system, where fruit is the secondary production. It was evaluated the variability in cone production as a function of the tree’s diameter at breast height and crown diameter and the individual tree’s competition status. The results indicate that stands managed in agroforestry systems with lower competition and individuals with larger diameter at breast height and crown diameter tend to produce more and heavier cones per tree. The first two principal components of the principal component analysis explain 84 % of the variance in cone production, trees’ dimensions and competition index. Tree competition status has a negative impact on production per tree.