退化森林生态系统恢复评价研究综述

森林退化是一个世界性的问题,对退化的森林进行恢复评价是合理地进行森林生态系统管理的基础。介绍了森林退化的概念,综述了退化森林生态系统恢复评价的一般程序,主要包括恢复目标的确定、参照系的选择、评价指标体系的构建及定量评价等几个方面。目前,大多数退化森林恢复评价主要包括物种多样性、植被结构和生态学过程3个方面。其中,物种多样性包括物种丰富度和多度等;植被结构包括植被盖度、乔木密度、高度、胸高断面积、生物量和凋落物结构等;生态学过程包括养分库、土壤有机质以及生物间的相互关系等。不同的研究者或管理者由于对恢复其生态系统服务功能的需求存在差异,评价退化生态系统恢复的角度也不一样。恢复评价可以从特殊种群到整个生态系统的不同层次进行。在深刻理解森林退化定义的基础上,建立现实的目标和正确地选择参照系是恢复评价的前提。     

茂兰退化喀斯特森林植被自然恢复中生态系统碳吸存特征

为了了解退化喀斯特森林自然恢复中生态系统碳吸存趋势, 采用空间代替时间的方法, 研究了茂兰退化喀斯特森林自然恢复中生态系统碳吸存特征。结果表明: 总体上植被生物量随恢复进程递增, 其中乔木层与其变化一致, 草本层、灌木层则相反; 喀斯特植被的地上与地下生物量之比较低, 尤其灌木层的地上与地下生物量之比最低; 加权平均含碳率随恢复进展递增; 随恢复进程, 植被乔木层碳密度递增, 草本层、灌木层碳密度递减; 总体上生态系统及其植被、土壤的碳密度由恢复早期(草本阶段、草灌阶段)经中期(灌木阶段、灌乔阶段)至后期(乔木阶段、顶极阶段)呈增加趋势, 而凋落物的相反。在贵州茂兰国家级自然保护区喀斯特森林的恢复进程中, 植被对生态系统碳库的影响最大, 尤其是木本植被, 而土壤的影响较小, 因此, 加强植被恢复对喀斯特地区生态系统碳汇具有极重要的意义。     

澳门九澳山海滨群落特征分析与生态恢复建议

采用群落生态学的方法研究了澳门九澳山海滨经人工抚育后恢复的植物群落特征。通过对其群落种类组成及多样性、空间结构和优势种年龄结构分析表明:1200 m2样地内共有维管束植物44科76属82种;藤本及中、小、矮高位芽植物占优势(86.59%),而大高位芽植物缺乏;群落优势种为豺皮樟(Litsea rotundifolia var.oblongifoli)、亮叶柃(Eurya nitida)、鸭脚木(Schefflera heptaphylla)和假苹婆(Sterculia lanceolata),均为增长型种群;九澳山海滨经人工抚育后恢复的植物群落植株高度较低。九澳山海滨土壤为赤红壤,其植被恢复策略可为赤红壤海滨的植被生态恢复提供参考。为促进海滨地带性植被的建设,我们还选择出海滨适生乡土树种29种,构建了多层次多树种的空间结构,并建议在九澳山海滨现有植被群落的基础上适当引种乔木树种、增加草本层植物种类。     

青藏高原东缘高寒森林小型兽类地表洞穴选择与生境特征

高寒森林地表的小型兽类洞穴是生态系统重要的景观特征,对于认识小型兽类的分布格局及其生态功能具有重要意义.为此,以青藏高原东缘典型高寒森林生态系统小型兽类洞穴为对象,于2015年夏季(7—9月)研究了其地表分布与生境特征.结果表明: 小型兽类洞穴平均洞口密度为182.8个·hm^-2,不同森林植被类型显著影响洞穴密度,表现为针叶林(328.6个·hm^-2)>针阔混交林(160.0个·hm^-2)>灌木林(125.0个·hm^-2)>阔叶林(81.5个·hm^-2)>竹林(66.7个·hm^-2). Mann-Whitney U Test和主成分分析表明,小型兽类栖息样地的乔木郁闭度、乔木均高、乔木胸径、乔木密度、地表植被盖度、倒木密度、倒木覆盖度、灌木均高都显著大于非栖息样地;乔木因子是决定小型兽类生境选择的首要因子,贡献率最大(44.8%),其次是倒木因子(23.2%)和灌草丛因子(13.9%).说明高寒森林地表有明显的小型兽类洞穴分布,其分布格局主要受植被和倒木的影响.     

河北小五台山国家级自然保护区森林群落与环境的关系

植物与环境之间的关系是一个复杂的演变过程,运用数量生态学方法探讨森林群落的物种组成、种群的生态特征、不同植物群落与环境之间的关系,有助于保护该区森林群落的稳定性和生物多样性。根据148个森林群落样方数据,选用双向指示种分析(TWINSPAN)和典范对应分析(CCA)方法,对河北小五台山国家级自然保护区森林群落进行分类和排序研究。结果表明:(1)TWINSPAN将该区的森林群落分为20个类型;(2)CCA排序结果较好地反映出群落分布格局与环境梯度的关系,各个森林群落类型在前两轴分异明显,在11个环境因子中,海拔、坡位、凋落层厚度、土壤导电率、土壤温度、土壤湿度、土壤厚度和干扰程度这8个环境因子对森林群落的分布起较大的作用,影响森林群落的分布格局,形成不同的植被类型。(3)乔木层优势种的CCA二维排序图所揭示的环境梯度与群落类型的分布有很大的相似性;t值双序图阐明了海拔、凋落层厚度、土壤温度、干扰程度等环境因子对森林群落乔木层优势种有着重要影响。采用TWINSPAN分类与CCA排序的方法,较好地解释了森林群落与环境因子的关系,为小五台山地区森林生态系统的科学管理和保护提供了理论依据,研究结果也为同类地区森林生态系统研究及保护提供参考和借鉴。     

川西亚高山不同森林生态系统碳氮储量及其分配格局

森林采伐和恢复是影响森林碳氮储量的重要因素。以川西亚高山岷江冷杉原始林、粗枝云杉阔叶林、天然次生林和粗枝云杉人工林为研究对象,采用样地调查和生物量实测的方法,研究了不同森林生态系统各组分碳、氮储量及其分配特征。结果表明岷江冷杉原始林、粗枝云杉阔叶林、天然次生林和粗枝云杉人工林生态系统碳储量分别为611.18、252.31、363.07 tC/hm~2和239.06 tC/hm~2;氮储量分别为16.44、12.11、15.48 tN/hm~2和8.92 tN/hm~2。恢复林分与原始林碳储量在土壤—植被的分配格局发生了变化,而氮储量未发生变化。岷江冷杉原始林以植被碳储量为主,恢复林分以土壤为主,氮储量均以土壤为主。乔木层碳储量分别占生态系统总储量的56.65%、17.63%、13.57%和22.05%,土壤层(0—80 cm)分别占32.03%、69.87%、76.20%和72.12%;土壤层氮储量占生态系统总储量的76.80%—92.58%。植物残体碳氮储量分别占生态系统总储量的4.40%—9.83%和2.94%—7.08%,林下植被所占比例最小。空间格局上,岷江冷杉原始林植被部分具有较高的碳储量,应进行保护。3种恢复林分具有较高的碳汇潜力,且地上/地下碳储量较低,表明其碳汇潜力尤其表现在地上部分。天然次生林利于土壤有机碳的积累,而人工林乔木层碳储量较高。     

林火干扰对广东木荷林生态系统碳库的影响

森林碳库在调节CO₂浓度及减缓温室效应中发挥重要作用。选择广东木荷林为研究对象,通过相邻样地法,进行植被生物量、凋落物生物量和土壤样品的采样与分析,研究不同林火干扰强度对生态系统各碳库(植被、凋落物和土壤有机碳)及生态系统碳库产生的变化规律和空间分布格局及其影响因素。结果表明:(1)植被碳密度随着林火干扰强度增强而减少,但不同组分的植被碳密度表现不同,乔木碳密度在不同林火干扰强度下变化与植被碳密度变化一致,而草本碳密度则呈现相反的变化趋势。相同林火干扰强度下,植被各组分碳密度均以乔木层降低幅度最大。林火干扰均显著降低了凋落物碳密度(P<0.05),并随林火干扰强度的增加其降低幅度增大,但不同林火干扰强度对凋落物碳密度的影响有所差异。林火干扰降低了土壤有机碳密度,且降低幅度随土层深度增加而逐渐变小。(2)林火干扰有效改变了生态系统碳库的空间分布格局。对照样地木荷林土壤有机碳库占比为61.59%,重度林火干扰后,土壤有机碳库占比为70.96%呈上升趋势,占生态系统碳库的优势地位,而植被和凋落物碳库占比呈下降趋势,处于生态系统碳库的次要地位。(3)双因素方差分析表明,林火干扰强度和土层深度及其交互作用均对土壤有机碳密度有显著影响。林火干扰强度解释了土壤有机碳密度变异的8.78%,土层深度解释了土壤有机碳密度变异的70.29%,林火干扰强度和土层深度之间的交互作用解释了土壤有机碳密度变异的8.16%。研究发现:林火干扰降低了生态系统碳库,且随林火干扰强度增加,生态系统碳库减少幅度增大。轻度林火干扰对森林生态系统碳库的影响差异不显著,而中度和重度林火干扰对森林生态系统碳库的影响差异显著。研究结果对深化亚热带森林固碳效应的影响机制提供理论支撑。     

内蒙古草甸草原地下芽库对极端干旱的响应

全球变化引发的极端气候事件严重影响草地生态系统结构与功能。然而,作为多年生草地重要繁殖体库,半干旱草甸草原地下芽库如何响应极端干旱尚未明确。本研究以内蒙古呼伦贝尔草甸草原为对象,利用模拟极端干旱样地平台,探究地下芽库及其与地上植被关系对模拟生长季极端干旱的响应。结果表明,环境降水水平与干旱处理下总芽密度分别为1443和1128芽·m^-2;分株总密度分别为1791和1346株·m^-2;群落整体分生组织制约系数分别为0.84和0.83。极端干旱对草甸草原地下芽库、地上分株密度以及二者之间关联均无显著影响。就优势植物功能群根茎型禾草而言,极端干旱对其地下芽库同样无显著影响。因此,地下芽库表现出对极端干旱一定程度的抵抗力,可作为草甸草原植被应对极端气候事件的保险策略,有效促进草甸草原在极端气候胁迫下的种群更新与植被恢复,有助于草甸草原植被稳定性与生态系统功能维持。     

森林植被恢复与环境生态因子互作关系研究进展

植被恢复是遏制生态环境恶化、改善脆弱生态系统的有效措施。植被同环境之间的关系极为复杂, 森林植被群落的恢复与重建, 在一定程度上改善了原有生境条件, 而不断被改变的环境生态因子反过来又影响着森林植被群落类型的演变。从土壤、群落、小气候3个方面综述了森林植被恢复与环境生态因子的互作关系, 针对现存问题, 认为今后应将微观的机理研究与宏观的动态研究相结合, 从不同尺度增强对植被退化、土壤、群落、小气候环境等因子互作关系的内在调控机理研究。     

不同区域森林火灾对生态因子的响应及其概率模型

火灾是影响森林生态系统过程的重要干扰之一,其对森林生态系统内各生态因子的响应各不相同.由于植被状况及生态环境的不同,森林火灾的时空分布特征在中国不同植被气候类型内表现不同,根据植被气候类型分类系统,将中国主要森林火灾地区划分为4个区域:东北(冷温带松林)、华北(落叶阔叶林)、东南(常绿阔叶林)和西南(热带雨林),应用遥感监测数据和地面环境数据,以时空变量、生态因子(植被生长变化指数、湿度等)为可选自变量,应用半参数化Logistic回归模型,就森林火险对不同生态影响因子的响应规律进行了分析,建立了基于生态因子的着火概率模型和大火蔓延概率模型,通过模拟及实际数据散点图、火险概率图,评估了模型应用价值.结果表明,土壤湿度及植被含水量在落叶阔叶林、常绿阔叶林、热带雨林地区对着火概率影响显著.在4个植被气候区内,土壤及凋落物湿度对大火蔓延的作用较小.在冷温带松林、落叶阔叶林、常绿阔叶林地区,植被生长的年内变化对火灾发生的影响显著,在常绿阔叶林地区,年内植被生长变化对大火蔓延的作用较小.森林火险概率与各生态因子的相关关系主要呈现出非线性.不同植被气候区内,火险概率受不同生态因子组合的影响,这与不同区域的植被状况及生态环境不同有关.在不同植被气候类型,应用时空变量、生态因子建立半参数化logistic回归模型,进行着火概率和大火蔓延概率的模拟具有可行性和实际应用能力.为进一步分析森林生态系统与火灾之间的动态关系、展开生态系统火灾干扰研究提供了理论基础.     

Vegetation composition,structure and soil properties across coastal forest–barren ecotones

Coastal barrens support rare plant species but may be threatened by forest encroachment. We determined whether trees spread into coastal barren habitat from forest patches and assessed plant species composition and soil properties across the forest–barren ecotone. We quantified tree age and height, soil properties, and vascular plant, bryophyte and lichen species composition along transects perpendicular to the edges of tree patches within the forest–barren ecotone in coastal Nova Scotia. Randomization tests assessed whether the vegetation and environmental characteristics were significantly different in the transition zone compared to one or both adjoining ecosystems. We used ordination to examine trends in species composition across the ecotone and the relationship to environmental variables. Tree age and height decreased continuously from the forest towards the edge of the forest patches. There were also trends in vegetation composition and structure from the forest into the open barrens. Many species were most abundant within the transition zone, although not always significantly. Soil properties were relatively uniform across the ecotone. The structure and vegetation of the forest–barren ecotone suggests that forest patches act as nuclei for forest expansion on barrens with a typical successional pathway where coastal barren vegetation is gradually replaced by forest species. This encroachment may pose a threat to rare barrens communities. While landscape factors such as salt spray and wind exposure may determine the general locations where forest can establish, biotic processes of growth and dispersal appear to govern the fine-scale expansion of tree patches.     

中亚热带喀斯特常绿落叶阔叶林空间结构特征

喀斯特常绿落叶阔叶林是中国西南喀斯特地区非地带性顶极森林群落,该研究基于相邻木关系的角尺度、混交度和大小比数3个结构参数,对木论国家级自然保护区常绿落叶阔叶林25 hm^(2)大型固定监测样地整体、上木层、下木层及其优势物种的空间结构特征进行了分析,以期揭示其森林空间结构现状,为喀斯特植被恢复重建提供依据。结果表明:(1)样地群落整体为轻度聚集分布格局,接近随机分布,物种高度混交且个体间大小分化程度相近,上木层呈现随机分布、强度混交和中庸偏亚优势状态,下木层呈现聚集分布、强度混交和中庸偏劣势状态。(2)样地内优势物种大多处于轻度聚集分布、强度混交且中庸偏劣势的状态。(3)林木大小比数与胸径、树高均呈显著负相关关系,混交度与胸径、树高均呈显著正相关关系,随林木的生长发育,树种优势度逐渐增大,物种多样性逐渐增强。研究认为,木论常绿落叶阔叶林空间结构尚未完全稳定,正处于演替中后期,具有发育成顶极群落的潜力。     

Tree–Shrub Interactions During Early Secondary Forest Succession in Uganda

Abstract One of the greatest challenges for ecologists this century will be restoring forests on degraded tropical lands. This restoration will require understanding complex processes that shape successional pathways, including interactions between trees and other plants. Shrub species often quickly invade disturbed tropical lands, yet little is known about whether they facilitate or inhibit subsequent tree recruitment and growth. We examined how shrubs and other vegetation (e.g., vines, grasses, herbs) affect tree recruitment, survival, and growth during the first 6 years of forest succession in Kibale National Park, Uganda. The study was undertaken in two recently logged exotic softwood plantations. We studied the successional trajectories in two recently logged areas that varied in their initial densities of trees and shrubs. Analyses suggested tree seedling presence and density were not strongly related to shrub density or height during succession. Tree sapling presence and density were positively significantly related to shrub density and height. We found little response in the tree community to experimental shrub removal, and although removal of all nontree vegetation temporarily enhanced tree growth, the effect disappeared after 2 years. Some early‐successional trees benefited from reduced competition, whereas some mid‐successional trees benefited from the presence of other vegetation. Some specific tree species responded strongly to vegetation removal. We interpret our findings in light of designing manipulations promoting forest restoration for biodiversity conservation and conclude with four tentative guidelines: (1) manage at the species level, not the community level; (2) increase facilitation for seedlings, reduce competition for saplings; (3) be cautious of assumptions about plant interactions; and (4) be adaptable and creative with new strategies when manipulations fail.     

基于无人机激光雷达遥感的亚热带常绿阔叶林群落垂直结构分析

森林植被高度与树木分布格局是植物群落重要结构特征,也是计算森林生物量分布的重要参数。传统的森林群落调查方法耗费大量人力物力难以进行较大尺度的群落结构测量,而一般的遥感影像也难以获得精确的地形信息及垂直结构。近年来激光雷达(Light Detection and Ranging,LiDAR)技术快速发展,能够较好的进行植被三维特征的提取并被广泛应用于森林生态系统检测模拟。且随着无人机低空摄影技术的发展催生的无人机激光雷达(UAV-Lidar)更增加了激光雷达的灵活性以及获取较大范围植被冠层信息的能力。而受限于激光的穿透性以及不同植被类型郁闭度的影响,该技术的应用多局限于在针叶林群落的垂直结构研究,而在常绿阔叶林的研究中应用较少。为探究现有无人机激光雷达设备及垂直结构提取分析技术应用于常绿阔叶林的可行性,利用无人机载激光雷达遥感技术对哀牢山中山湿性常绿阔叶林3块面积1hm~2的样地进行基于数字表面模型以及数字地表高程模型做差得到树冠高度模型测量的植被冠层高度、基于局部最大值法进行单木位置提取并使用Clark-Evans最近邻体分析方法进行样地内高大乔木分布格局的计算。分析结果显示,植被高度提取精度平均大于95%,与地表实测的植被高度值拟合度较高,相关系数R~2介于0.833—0.927之间;3个样地冠层高度平均值分别为18.79、19.08、17.03 m,标准差分别为8.10、7.34、7.17 m。单木探测百分比平均86.3%,用户精度以及生产者精度平均分别为75.69%和65.15%。实测得出三个样地全部高大乔木空间分布格局均为聚集分布,而激光雷达测量结果显示为随机分布或均匀分布。实验显示基于无人机激光雷达技术能够很好地提取植被冠层高度信息并能够较好地获取树木位置,但对于树木空间分布格局判定的准确性有待于进一步探索。未来研究应从多角度对激光雷达测量造成的误差原因予以分析(如环境因素),并进一步研究更为精确的单木提取以及植被高度提取方法,为通过无人机激光雷达测算森林生物量及各种生态过程提供更加精准的指标数据。     

Nurse Trees as a Forest Restoration Tool for Mixed Plantations: Effects on Competing Vegetation and Performance in Target Tree Species

Multi‐species mixed plantations can be designed to meet social, economic, and environmental objectives during forest restoration. This paper reports results from an experiment in southern Sweden concerning the influence of three different fast growing nurse tree species on the cover of herbaceous vegetation and on the performance of several target tree species. After 10 years, the nurse trees had reduced the competing herbaceous vegetation but the effect was weak and it may take more than a decade to achieve effective vegetation control. The nurse tree species Betula pendula and Larix x eurolepis did improve stem form in some target tree species, but had a minor effect on survival and growth. The open conditions before crown closure of nurse trees strongly influence seedling performance and so delayed planting of target tree species may provide a means to avoid those conditions. Survival and growth differed greatly among the tree species. Besides the two nurse tree species mentioned above, high survival was found in Picea abies and Quercus robur and intermediate survival in Fagus sylvatica, Tilia cordata, and in the N‐fixing nurse tree Alnus glutinosa. Survival was low in the target tree species Fraxinus excelsior L. and Prunus avium. For restoration practitioners, our results illustrate the potential of using nurse trees for rapidly building a new forest structure and simultaneously increase productivity, which might be a cost‐effective strategy for forest restoration.     

沿海基干林带结构调控对林分冠层结构参数及林地土壤的影响

以上海浦东沿海水杉基干林带为研究对象,分析水杉林分间伐以及间伐后林下套种其他树种构建复层林2种结构调控措施下林分冠层及林地土壤的变化。结果表明:间伐和复层林构建可显著促进上层乔木树高和胸径的生长,也有助于林下乔灌木和草本植物的发育;未间伐水杉林带LAI最高,其次为复层林带,间伐林分最低。各林分林隙分数的分布特征与LAI相反;3种林分土壤全N、全P和速效K含量差异不显著,而间伐林和复层林土壤速效P、水解N和有机质含量比未间伐林有进一步改善,但间伐林分和复层林带之间土壤养分状况没有显著差异;林分结构调控增加了0～50cm土壤有机碳储量,其中以0～15cm土层有机碳密度最高。复层林带和间伐林土壤呼吸速率比未间伐林分别提高了42%和33%。     

黄土高原植被-降水关系的临界现象及其在植被建设中的意义

以退耕还林还草为主要内容的大规模的生态环境建设正在我国兴起,如何进行科学的植被配置规划,是一个急待解决的理论和实践问题。以自然植被初级生产力(NPP)表征潜在的即最大可能的植被条件,并与现在的森林覆盖率分布以及植被构成特征进行了比较。通过大量实测资料的分析,查明了降水条件对森林覆盖率和NPP的影响及其临界条件。研究表明,以NPP表征的天然植被特征值在黄土高原地区的分布具有显著的地带性。NPP与年平均降水量的关系中存在着两个临界点,分别代表着植被类型的变化,Pm=250mm可视为荒漠与草原植被之间的临界点,而Pm=480mm则为草原(森林草原)与森林(落叶阔叶林)之间的临界点。尽管次生的人工植被受人为因素的控制,由于存在着自然稀疏化过程,将使人工林地最终达到某种与水分承载能力相一致的平衡状态,自然条件如年降水仍然是决定其覆盖度的重要因素。在森林覆盖率与年平均降水量的关系中表现出一个临界点。当Pm<480mm时,森林覆盖率很低,且随年降水的增大而增大的速率很慢;当Pm>480mm时,森林覆盖率随年降水量而急剧增大。人为建造的水土保持植被,是一个自然的人工生态系统,也是一个自组织系统,它通过自我调节而趋向于达到某种平衡,建立一定的群落结构甚至于趋向于某种顶极群落,从而实现乔木、灌木、草本植物组成的最优组合。从这一原理出发,提出了一个基于实测资料进行统计分析的方法,来确定使乔木面积在乔木、灌木、草本总面积中的最优或较优的比例,得出了具体的数据。     

Aspect and slope as determinants of vegetation composition and soil properties in coastal forest backdunes of Eastern Cape,South Africa

Coastal sand dune ecosystems are subjected to many stress and disturbance factors that are particularly high in the foredunes compared to the backdunes. Although a few studies have been conducted on eastern coastline sand dunes of South Africa, none have examined the relationship between aspect and slope on vegetation composition and soil properties of coastal forest backdunes. Vegetation and soil sampling were conducted in 11 transects, each with four plots measuring 10 × 10 m, located on the seaward and landward sides and on middle and lower slopes of backdunes of Bathurst coastal forest. A total of 39 species were identified, of which 23 were trees and shrubs, thirteen were forbs and three were grasses. The data show that both aspect and slope had limited influence on vegetation community assemblage and soil properties, but had significant effects on individual species distribution. There was a grass‐dominated community on the middle slope and a tree‐ and shrub‐ dominated community on the lower slope. These two plant communities act as the required coastal forest ecosystem engineer driving variability in soil properties between the slopes, the most prominent being high soil nutrients and moisture in the lower slopes compared to the middle slopes.     

Water availability as driver of birch mortality in Hustai National Park,Mongolia

Trees in the forest-steppe ecotones face stress due to reduced water availability as a consequence of more extreme seasonal fluctuations in precipitation and temperature. Together with browsing pressure this can hinder tree growth, tree regeneration and competition between trees and grasses. We studied the impact of both stress factors on the mortality of birch trees in two forest sites at Hustai National Park, Mongolia, by applying tree-ring research to determine growth-limiting factors and assessing browsing pressure on young and adult birch. We expected warm and dry summer conditions as main growth limiting factor. Moreover, we expected a positive relation between deer density and tree mortality with browsing mainly affecting smaller trees with a low diameter at breast height (DBH). We found that the growth in both birch populations is mainly driven by winter precipitation and – to a lesser extent – negatively affected by high summer temperature. This suggests that water availability as defined by soil moisture, especially at the beginning of the growing season is crucial for birch growth in our study area. For mortality we found significant differences between both populations, but no significant relationship with deer density. In plots with high mortality rates mean tree height of the remaining living trees decreased. These results suggest that under expected climate change with declining annual precipitation rates, the birch forest of Hustai National Park is converting into a steppe ecosystem, like described for other forest ecosystems in this ecotone.