Simulation modeling of the effects of site conditions and disturbance history on a boreal forest landscape

Abstract. Computer simulations were used to elaborate hypotheses about controls on forest structure and composition in a 0.7 km ≤ area of boreal forest in Central Sweden. DBH and species of all adult trees and stand conditions were recorded for 57–10 m radius plots. Ordination of these data suggested that nutrient-availability and time-since-disturbance were the main controls of forest composition and structure within the area. The simulation model couples equations representing the effect of tree canopy structure and biomass on light and soil conditions with equations representing the effect of these conditions on reproduction, growth and mortality in height cohorts of trees on a 0.1 ha patch. Nitrogen-availability levels for each modeled plot were simulated by species-specific growth multipliers. The model was run for 400 simulated yr at six levels of N availability. Age and N status of each study plot were inferred by matching with the most similar model output. Inferred ages agreed with what is known of the disturbance history, and site factors related to soil fertility were correctly correlated with the inferred N status. The consequences of size-selective disturbance were explored by model experiments. Biomass was removed from large or small size classes at 100 - 200 yr and the simulations were run for an additional 300 yr. Disturbed stands of high N status often became similar to undisturbed stands of different N status. Size-selective disturbances produced stands that were different from any in the undisturbed succession, but these differences disappeared within 50 - 100 yr, implying successional convergence in stand structure and composition. Plots of simulated basal area against time and nitrogen-availability for the four species illustrate the time dependence of species performance along a fertility gradient.     

A role-type model (rope) and its application in assessing climate change impacts on forest landscapes

Gap-phase replacement is a general phenomenon found in forest ecosystems, worldwide. Different tree species can be expected to produce different sizes of gaps when they die. Species also vary in their regeneration success in gaps of different sizes. In this paper, the gap-phase interactions among tree species in a forest stand are simulated by a role-type stand model called ROPE. By incorporation of environmental effects on tree height, ROPE can simulate forest composition and stand leaf area under different climate conditions. The model was developed for forest ecosystems in northeastern China and was used to simulate the forest landscape structures under current climate conditions and under four climate change scenarios for greenhouse gas related warming. These scenarios were obtained from general circulation models developed by different atmospheric research centers. Korean pinebroadleaf mixed forest and larch forest are the major stand types in the study area under present conditions. Under the four climate change scenarios, Korean pine-broadleaf mixed forest would be expected to occur only on the higher parts of large mountains. Larch forest only would be found north of the study area. Broadleaf forest would become the dominant vegetation over the study area. Use of the Kappa statistic to test for similarity in spatial maps, indicates that each climate change scenario would result in a significant change of forest distributions.Supported by The United States National Science Foundation Grant BSR-8702333 to University of Virginia.     

林窗模型的几个基本问题的研究:I.模拟样地面积的效应

以中国东北长白山阔叶红松林为例,应用林窗模型NEWCOP探索了不同模拟样地面积对林窗模型输出结果的影响.结果表明,模拟样地面积大小变化可影响模拟出的森林群落的树种组成和模拟样地的林窗出现周期,通过应用这一特点确定了阔叶红松林的林窗面积为400～800m².     

中国东北森林生长演替模拟模型及其在全球变化研究中的应用

 NEWCOP模型是一个新的适于模拟东北森林的种类组成动态的林窗类计算机模拟模型,它通过模拟在每一个林分斑块上的每株树木的更新、生长和死亡的全过程来反映森林群落的中长期生长和演替动态。由于 NEWCOP模型是一个由气候变量驱动的生态系统模型,故可用于评价气候变化对东北森林生长和演替的影响。在东北大兴安岭、小兴安岭和长白山地区对NEWCOP模型进行了验证和校准。沿环境梯度对NEWCOP模型的数字模拟实验表明：它能准确地再现顶极森林中树种组成及其在东北地区的垂直分布规律和水平分布规律;能准确地再现大兴安岭、小兴安岭和长白山的主要类型森林的生长和演替规律;在一定的场合NEWCOP还可反映林分的径级结构;NEWCOP模型还具有对现有森林的跟踪模拟能力。应用NEWCOP模型评估了东北森林生态系统对可能气候变化的敏感性。在GFDL 2×CO2和GISS 2×CO2气候变化情景下,东北森林的种类组成将发生很大变化,落叶阔叶树将取代目前长白山、小兴安岭的红松(Pinus koraiensis)和大兴安岭的兴安落叶松(Larix gmelinii)成为东北森林主要树种,而针叶树将在地带性森林中占很小的比重,阔叶树中蒙古栎(Quercus mongolica)将是最重要的树种,它将成为小兴安岭和大兴安岭最主要树种;东北地区适于森林生长的区域将大幅度减少,这些变化主要发生在气候变化过渡期。东北森林对不同的气候变化情景有不同响应。但是,总的趋势是未来东北森林中落叶阔叶树的比重将大幅度增加。这些结论对在全球气候变化背景下,我国东北合理地选择造林树种和制定现有森林的保护经营策略具有一定参考价值。     

Tree community features of two stands of riverine forest under different flooding regimes in Southern Brazil

Two 1 ha plots of a Southern Brazilian subtropical riverine forest, subject to different frequency and duration of floods, were compared to detect the differences in physiognomic structure, tree community composition, richness and diversity. Each plot was made up of 100 contiguous 10×10 m subplots, where 3451 trees with pbh 15 cm were measured and identified. The survey observed 30 tree species, in the frequently flooded plot and 48 in the occasionally flooded plot. A detailed topographical and soil survey was carried out in both plots and indicated that the levels of organic matter and most mineral nutrients were higher in the frequently flooded stand. The forest understory was denser in the occasionally flooded stand which also showed taller emergent trees. Multivariate ordination and grouping techniques showed that the species’ abundance distribution was strongly related to the topographical variation. There was a clear pattern of species turnover according to topographic position, indicating that tree species developed different abilities to survive flooding events. As a result, the two plots also differed in their tree frequency per species regeneration, vertical distribution and dispersion groups. Both species richness and diversity decreased with increasing flood frequency, also showing a patchy distribution within both stands. At a local scale, flooding regime is regulating the spatial variation of α-diversity by forming different seral stages of predictable species composition. Compared to regularly flooded riverine and floodplain forests, riverine forests, with unpredictable flooding regimes, may show higher diversity at a local scale and more abundant opportunistic species of high environmental plasticity.     

A GIS-based simulation program to predict multi-species size-structure dynamics for natural forests in Hokkaido, northern Japan

A simulation program that runs on a geographic information system (GIS) was developed to predict the multi-species size-structure dynamics of forest stands. Because important characteristics of a forest stand, including woody biomass accumulation, carbon storage, commercial value of timber, and functions for environmental conservation, can be inferred from the size structures of the component populations, management plans can be made from the predictions of the size-structure dynamics. For example, the simulation can incorporate various forms of thinning; forest managers can then try several thinning plans in simulated forest stands and choose the appropriate plan that achieves the best results. Using GIS to predict the size-structure dynamics of forest stands is of practical importance, because GIS has been used widely in forest management and can easily handle spatial distributions of environmental information (e.g., climate, geology, soils) that may influence tree performance. To predict size-structure dynamics, the program numerically solves a continuum equation that describes size-structure dynamics based on growth and mortality rates of individual trees. When predicting size-structure dynamics of a forest stand, the program obtains the environmental information of the stand from a database stored in the GIS and calculates environmental factors such as warmth index and potential evapotranspiration/precipitation ratio that influence growth and mortality rates. The simulation program calculates growth and mortality rates using published growth and mortality models that incorporate the effects of size of the individual, competition between trees, and abiotic environmental factors. To demonstrate the effects of abiotic environmental factors on the multi-species size-structure dynamics, sensitivity analyses were conducted. The size-structure dynamics varied in a way that was predictable from the responses of the growth and mortality rates to variations in the abiotic environmental factors. To demonstrate the size-structure dynamics in different locations, five test runs of the simulation program were also performed using the same initial size-structure and five different sets of abiotic environmental conditions from five locations. At the end of the simulation, the predicted size structures differed because the growth and mortality rates differed among the five locations. Finally, the response of the size structure to thinning was clarified. The result showed how the size structure of a component species in a forest stand is dependent on the presence of other species.     

Spatial pattern and process in forest stands within the Virginia piedmont

Abstract. Question: Underlying ecological processes have often been inferred from the analysis of spatial patterns in ecosystems. Using an individual‐based model, we evaluate whether basic assumptions of species’life‐history, drought‐susceptibility, and shade tolerance generate dynamics that replicate patterns between and within forest stands. Location: Virginia piedmont, USA. Method: Model verification examines the transition in forest composition and stand structure between mesic, intermediate and xeric sites. At each site, tree location, diameter, and status were recorded in square plots ranging from 0.25 to 1.0 ha. Model validation examines the simulated spatial pattern of individual trees at scales of 1–25 m within each forest site using a univariate Ripley's K function. Results: 7512 live and dead trees were surveyed across all sites. All sites exhibit a consistent, significant shift in pattern for live trees by size, progressing from a clumped understorey (trees ± 0.1 m in diameter) to a uniform overstorey (trees > 0.25 m). Simulation results reflect not only the general shift in pattern of trees at appropriate scales within sites, but also the general transition in species composition and stand structure between sites. Conclusions: This shift has been observed in other forest ecosystems and interpreted as a result of competition; however, this hypothesis has seldom been evaluated using simulation models. These results support the hypothesis that forest pattern in the Virginia piedmont results from competition involving species’life‐history attributes driven by soil moisture availability between sites and light availability within sites.     

森林动态模型概论

讨论了模拟森林林分动态变化的模型,并把模型分为森林生长模型和演替模型。森林生长模型包括：全林分模型、林分级模型和单木模型;演替模型包括马尔可夫类模型和林窗模型。文中给出了演替模型的基本原理和适用性,在比较早期和最新发展的林窗模型后,叙述了林窗模型的新进展。生长和演替模型的结构和数据要求不同决定了它们的在时间和空间上的适应性,最后指出模型将向综合总体方向发展     

目标树抚育对亚热带天然次生灌丛群落结构和多样性的影响

在浙江省临安市选择典型天然次生灌丛,分别进行封禁和目标树抚育,探讨灌丛恢复为乔木林的可能性.结果表明: 4年后,与未管护(对照)相比,封禁和目标树抚育后群落平均胸径分别提高1.3和2.6倍,平均高度分别提高0.5和1.1倍;目标树抚育林木出现了对照林分没有的4.5～8.5 cm径阶和4.5～8.5 m树高阶,形成了4 m高的新林层;灌木层物种丰富度和多样性指数没有因抚育而下降;封禁管理维持了群落的树种组成,遵循原有的演替方向;目标树抚育显著改变了群落的树种组成,提高了目的树种的重要值,近期有可能恢复成为针阔混交林群落.与封禁相比,目标树抚育在优势林木胸径和高度生长、树种组成改善等方面更能达到预想的目标.在有条件经营的情况下,可以选择目标树抚育模式对天然次生灌丛进行管理,从而达到加快群落恢复演替形成乔木林的目的.     

Compensatory mechanisms of central U.S. hardwood forest communities in a changing environment

Although it is known that forest productivity is highly correlated with canopy leaf area, the influence of environmental gradients within a forest on leaf area-productivity relationships has not been well-documented, particularly for mixed-species forests. The present study was conducted to define the mechanisms by which Central Hardwood forest communities respond to changes in environment. Ecological Land Type Phases (ELTPs) of the Central Hardwood region, delineated by soils, topography, and indicator vegetation, provided a framework to examine the impacts of environmental gradients on leaf area-productivity relationships at a landscape level. Three ELTPs- (1) Quercus prinus / Carex picta-Vaccinium, Dry slope, (2) Quercus alba-Acer saccharum / Parthenocissus, Dry-Mesic Slope, and (3) Fagus-Acer saccharum / Arisaema, Mesic Slope- were selected and stratified into three slope positions viz. the upper edge of the community, the middle or center of the community on the slope, and the lower edge of the community. Circular plots of 0.02 ha in size were established for sampling. Standing stock (basal area, volume, and biomass), annual production, leaf area index, and growth efficiency were determined for each slope position and examined to determine the mechanisms and trends for community change across slope positions. Among the three communities studied, dry slopes compensated for drier conditions through both a shift in species composition and reduced annual production. Dry-mesic slopes exhibited adaptation only through changing species composition whereas mesic slopes compensated for drier conditions primarily through reduction in annual production. These results indicate that mixed-species forest communities adjust to environmental gradients through several mechanisms and that the relative importance of these mechanisms changes with community type.     

Vegetation patterns and environmental gradients in tropical dry forests of the northern Yucatan Peninsula

Patterns of plant species composition and their relationships to soil and topographic variables were investigated in tropical dry forests across the north central Yucatan, Mexico. Seven sites were studied in the oldest accessible forests along a 200–km transect oriented northwest to southeast; an eighth site was located in a little‐disturbed area located 75 km northeast of the transect. Two of the sites were on Mayan ruins. All sites were sampled using 9–24, 10m × 20m plots (<n= 132) for woody stems ≥ 3.0 cm diameter breast height. The important natural forest species were Bursera simaruba, Caesalpinia gaumeri, Gymnopodium floribundum, Piscidia piscipula, and Thouinia paucidentata. The two most important woody species in ruin woodlands were Brosimum alicastrum and Croton lundellii. Forest plots (n=108) had 17 species on average, ruin plots (n= 24) nine species. Mean basal area of stems at the forest plots (20.7 m².ha^‐1) was lower than in ruin plots (28.4 m².ha^‐1). Detrended Correspondence Analysis generally placed plots by site along the geographic transect. Natural forest plots and sites were separated from the plots on ruin sites. The five soil and topographic variables (slope, soil depth, percent surface rock, soil pH, total soil organic matter) differed significantly among sites. Plot values were correlated with DCA axe scores. Intersite floristic variation reflects an overall west to east environmental gradient affected by climate.     

Land-use history affects understorey plant species distributions in a large temperate-forest complex,Denmark

In Europe, forests have been strongly influenced by human land-use for millennia. Here, we studied the importance of anthropogenic historical factors as determinants of understorey species distributions in a 967 ha Danish forest complex using 156 randomly placed 100-m² plots, 15 environmental, 9 spatial, and 5 historical variables, and principal components analysis (PCA), redundancy analysis (RDA) as well as indicator species analysis. The historical variables were status as ancient (1805 AD) high forest, reclaimed bogs, ≤100 m from Bronze Age burial mounds, or former conifer plantation, and stand age. The PCA results showed that the main gradients in species composition were strongly related to the explanatory variables. Forward variable selection and variation partitioning using RDA showed that although modern environment was the dominant driver of species composition, anthropogenic historical factors were also important. The pure historical variation fraction constituted 13% of the variation explained. The RDA results showed that ancient-forest status and, secondarily, reclaimed bog status were the only significant historical variables. Many typical forest interior species, with poor dispersal and a strong literature record as ancient-forest species, were still concentrated in areas that were high forest in 1805. Among the younger forests, there were clear floristic differences between those on reclaimed bogs and those not. Apparently remnant populations of wet-soil plants were still present in the reclaimed bog areas. Our results emphasize the importance of historical factors for understanding modern vegetation patterns in forested landscapes.     

EVALUATING THE IMPACT OF INSECT COMMUNITY ON PINE CATERPILLAR DENSITY IN DIFFERENT STAND CONDITIONS

Abstract  Natural enemies play an important role in inhibiting population growth of pine caterpillar, but they are difficult to be used in systematic management models. This paper attempts to use some forest stand factors to substitute the effect of natural enemies, which could be used in management models. Through the studies from more than 200 stand plots with different stand conditions in Qu County, Changshan County, and Longyou County of Zhejiang Province, 16 predominant species groups from insect community were selected, and the canonical correlation coefficient between the diversity index, species number, individual number of predominant species and the stand factors was analyzed. The results indicated that the canopy density and ground vegetation were the key stand factors for characterizing the function of insect community. The forest area was divided into four types according to the variation of canopy and vegetation conditions by using cluster analysis. The regression models between the rate of variation of pine caterpillar density and the forest factors including canopy density and vegetation covers were developed in each type of forests. There were significant differences in diversity index, species number, individual number of insect community and population density of pine caterpillar among four types of forests, and correlation coefficients of the four regression models were rather high ( r =0. 924 — 0. 964). This means that the classification of the forest stands is reasonable and the stand factors can characterize the effect of natural enemies in practice.     

Individual-based model of spatio-temporal dynamics of mixed forest stands

This research presents the results of constructing and parameterizing an individual-based model of spatiotemporal dynamics of mixed forest stands. The model facilitates computerized experiments with forest stands having different combinations of species and age structures. These forest stands grow on temperate areas where light is the main system-forming factor that shapes and develops forest ecosystems. The model TEMFORM (TEMperate FORests Model) is developed with few equations and parameters, most of which can be estimated using standard forest inventory data. Parameterization of the model used the growth tables of a set of basic forest-forming species in Far East Russia. Simulation results of the development of the natural single- and mixed-species stands and the effects of different types of disturbances on the stand dynamics and compositions are presented.     

Long‐term seed bank dynamics in a temperate forest under conversion from coppice‐with‐standards to high forest management

Questions: How do changes in forest management, i.e. in disturbance type and frequency, influence species diversity, abundance and composition of the seed bank? How does the relationship between seed bank and vegetation change? What are the implications for seed bank dynamics? Location: An ancient Quercus petraea — Carpinus betulus forest in conversion from coppice‐with‐standards to regular Quercus high forest near Montargis, France. Methods: Seed bank and vegetation were sampled in six replicated stand types, forming a chronosequence along the conversion pathway. The stand types represented mid‐successional stages of stands in transition from coppice‐with‐standards (to high forest (16 plots) and early‐ and mid‐successional high forest stands (32 plots). Results: Seed bank density and species richness decreased with time since last disturbance. Adjusting for seed density effects obscured species richness differences between stand types, but species of later seres were nested subsets of earlier seres, implying concomitant shifts in species richness and composition with time since disturbance. Later seres were characterized by species with low seed weight and high seed longevity. Seed banks of early seres were more similar to vegetation than to later seres. Conclusions: Abandonment of the coppice‐with‐standards regime altered the seed bank characteristics, as well as its relationship with vegetation. Longer management cycles under high forest yield impoverished seed banks. For their persistence, seed bank species will increasingly rely on management of permanently open areas in the forest landscape. Thus, revegetation at the beginning of new high‐forest cycles may increasingly depend on inflow from seed sources.     

基于线性分位数混合效应的辽东山区红松冠幅模型

冠幅是反映单木生长状态及构建林木生长收获模型的重要变量。本研究以辽东山区大边沟林场10～55年生红松人工林为对象,基于66块固定样地的2763株红松的每木检尺数据,选取冠幅基础模型,采用再参数化的方法引入单木竞争指标(R_d),利用哑变量的方法引入了林分密度、林层变量,构建不同分位点(0.50、0.90、0.93、0.95、0.96、0.99)的冠幅分位数回归模型,并与传统方法进行比较,选取模拟林分最大冠幅的最优分位点。为反映林分中单木冠幅在林木个体之间的差异,建立了基于样地水平的最优分位点的线性混合效应分位数回归冠幅模型,分析各变量对单木冠幅的影响。结果表明: 基于F统计检验,不同林分密度和林层的冠幅模型具有显著差异,在基础模型中引入林层、林分密度和竞争后,模型R_a²提高0.0104,均方根误差降低0.0115,均方误差降低为7.4%;与最小二乘法比较,分位数回归模型能够较好地模拟林分状态下的单木最大冠幅,并选出0.96分位点和0.93分位点作为上林层和下林层的分位数回归模型的最优分位点。引入混合效应的线性分位数回归模型的赤池信息准则、贝叶斯信息准则、HQ信息准则等评价指标优于传统分位数回归,参数标准误显著降低,混合效应的引入很好地解释了样地之间的差异。就上林层和下林层而言,林分密度越大,最大冠幅越小;相对直径越大,最大冠幅越大,其中林分密度对下林层的冠幅影响大于上林层,当林分密度足够大时,冠幅随着胸径的增大先增大后降低。本研究构建的基于混合效应的分位数回归模型能有效提高模型的拟合优度,今后可通过调控林分密度、适度抚育间伐等措施,实现对辽东山区红松人工林的科学营建和可持续发展。     

Mesoscale distribution of understorey plants in temperate forest (Kalø, Denmark): the importance of environment and dispersal

We studied the influence of environmental heterogeneity and dispersalprocesses on mesoscale distribution patterns of understorey plants in a 330ha ancient, managed temperate forest area. Similarity matrixregression analysis showed that overall species similarity was affected bysite(the two disjunct halves of the forest area), spatial autocorrelation at the100m scale, edaphic conditions, stand type and stand structure, andthe occurrence of open areas, but not by stand age or topography. Usingautologistic regression we analysed the influence of clumping, site, edaphicconditions, stand composition, open areas, and stand structure on theindividualdistribution patterns of the 60 most abundant species. Only five species haddistributions that were not significantly related to any of the explanatoryvariables. A large number of species showed significant, and often differing,relationships to at least one of the environmental parameters. At least 20species exhibited clumping at the 100 m scale that was independentof the environmental parameters. Principal coordinate analysis and an analysisusing the Ellenberg ecological indicator values suggested that no importantgradients had been overlooked. Dispersal ability (estimated from dispersalmode)and clumping independent of environmental heterogeneity were related. Poordispersers exhibited stronger clumping at the 100 m scale thangooddispersers. Our results support the joint importance of environmentalheterogeneity and dispersal processes in shaping the mesoscale distributionpatterns of forest herbs. We conclude that the distribution of plant species,and species coexistence and species composition in plant communities, cannot beexplained without simultaneously considering environmental heterogeneity anddispersal.     

基于结构方程模型的兴安落叶松天然林更新影响因素

以大兴安岭新林林业局翠岗林场49块兴安落叶松不同蓄积占比的固定样地调查数据为基础,分别从林分非空间结构、林分空间结构、林木多样性、土壤条件和立地条件5个方面37项观测指标,在相关性分析的基础上,构建林分尺度更新密度和更新多样性(包括树高和地径)的结构方程模型,并计算各路径的直接、间接和总影响系数,量化和提取影响大兴安岭地区兴安落叶松天然林更新密度和更新多样性的关键可控因素,为该地区森林的可持续经营提供理论基础和技术依据。结果表明: 对林分更新密度影响最大的潜变量依次为:林分非空间结构(-0.410)>林木多样性(0.380)>土壤条件(0.250)>立地条件(0.249)>林分空间结构(0.197);对林分更新多样性影响最大的潜变量依次为:土壤条件(0.778)>立地条件(0.748)>林分空间结构(0.684)>林分非空间结构(0.287)>林木多样性(0.105)。综合来看,影响更新密度和多样性的共性观测变量主要为:pH值、全钾、树种多样性、树高多样性、角尺度和单位蓄积。在后续经营中,可通过采伐或补植阔叶树种的方式来优化和调整林分的树种组成、多样性、土壤pH值和养分等特征,最终达到促进天然更新的目的。     

不同立地条件下昆虫群落对松毛虫密度影响的估计(英文)

天敌在松毛虫种群动态中起重要作用,但把天敌的作用加入系统管理模型中却很困难。本文试图用林分因子代替天敌的作用,因为林分条件可以方便地用于系统模型中。通过对浙江省衢县、常山县和龙游县不同立地条件的222块样地的调查数据的分析,从70多科的昆虫群落中选择了16个关键种（科）,并对林分因子和各样地关键种的种数、个体数、多样性之间的关系进行了典型相关分析。结果表明郁闭度和植被条件（灌木层盖度、灌木层高度、草木层盖度、草木层高度、总盖度）是刻划昆虫群落功能的关键林分因子。根据郁闭度的变化和植被条件用系统聚类方法可以把林区分为四种类型,这四种类型林地的昆虫多样性指数、种数、个体数量和松毛虫的密度都存在明显差异;在每种林地都建立了松毛虫密度的变化率与关键林分因子间的回归模型,所有的回归的相关系数都相当高（0．924－0．964）,因此可以认为对林地的分类是合理的,而且用林分因子评估天敌的影响在实践中是可行的。