贵州省普定县不同植被演替阶段的物种组成与群落结构特征

通过对贵州省普定县喀斯特地区不同植被演替阶段群落的调查, 研究了植被演替过程中群落物种组成和群落结构的变化。结果表明, 该地区的植被主要处于5个演替阶段, 即次生乔木林、乔灌过渡林、藤刺灌丛、稀灌草丛以及火烧干扰后的蕨类植物群落。本次调查共记录到植物365种, 隶属89科218属。其中, 蕨类植物31种, 隶属14科23属; 种子植物334种, 隶属75科195属。物种分布较多的科主要有蔷薇科、菊科、禾本科、百合科、忍冬科、唇形科、莎草科、樟科、葡萄科和水龙骨科。随着正向演替的推进, 物种丰富度增加, 群落结构趋于复杂化。藤刺灌丛与乔灌过渡林群落层次不明显, 次生乔木林分层明显。从藤刺灌丛向次生乔木林演替的过程中, 小径级个体所占比例明显降低, 高于1.3 m植物的总密度、乔木密度和藤本密度都先升高后降低, 而灌木密度呈逐渐降低的趋势。对喀斯特地区植被的恢复提出了参考措施。     

森林群落复杂性分析:以广东黑石顶森林为例

本文将森林群落复杂性定义为消除群落中任意一个树木个体的种名及层次不确定性所需的平均信息量,提出用联合熵H(X,Y)测度群落复杂性.H(X,Y)=H(X)+H(Y|X),其中H(X)=-∑Si=1(ni)/(N)log2((ni)/(N)),H(Y|X)=-∑Si=1(ni)/(N)∑4j=1(nij)/(ni)log2((nij)/(ni)),分别为树种组成复杂性和树种结构复杂性.式中S为森林群落树种数,N为森林群落的树木总株数,ni(i=1,2,...,S)为第i个树种的株数,nij(j=1,2,3,4)为第i个树种在第j层次的株数.用联合熵分析了广东省封开县黑石顶自然保护区针叶林、针阔混交林和常绿阔叶林等3种典型森林类型的复杂性.结果表明,群落复杂性的顺序为常绿阔叶林>针阔混交林>针叶林.同时,各森林类型的群落复杂性H(X,Y)与取样尺度之间具有较好的分形关系.     

广东黑石顶自然保护区森林次生演替过程中的群落动态

对黑石顶自然保护区森林次生演替过程中的群落结构、物种组成及物种多样性变化的研究结果表明：在皆伐裸地恢复阶段最初的２～４ 年,先锋种、阳生性种及中生性种的幼苗同时大量出现于次生裸地上;演替至１０ 年时,群落中先锋种的数量占绝对优势,阳生性种保持稳定,中生性种趋于减少;针阔叶混交林阶段,先锋种趋于减少,阳生性种趋于增加;阳生性常绿阔叶林阶段,先锋种基本衰退,而阳生性种占绝对优势,同时中生性种趋于增加;演替至中生性常绿阔叶林阶段,阳生性种逐渐消退,中生性种的数量占优势。皆伐裸地恢复阶段为群落垂直结构的形成期,仅有Ⅲ级以下立木,乔木层无明显分层,群落的个体密度无明显变化;由恢复阶段进入针阔叶混交林初期为群落垂直结构分化期,群落中出现大量Ⅲ、Ⅳ级立木,乔木层可分为３ 个亚层,但群落的个体密度因自疏作用而显著下降;针阔叶混交林阶段为群落垂直结构的相对稳定期,Ⅴ级立木多度和胸高断面积呈演替系列的第一个高峰,同时群落的个体密度显著增大;阳生性常绿阔叶林阶段为群落垂直结构的变动期,群落中Ⅴ级立木多度和胸高断面积大幅度下降,个体密度无明显变化;中生性常绿阔叶林阶段为群落垂直结构的稳定期,群落中各立木级趋于更加完善,胸高断面积达到演     

普定喀斯特地区不同演替阶段植物群落凋落物动态

凋落物是陆地生态系统碳收支和物质循环的重要组成部分.本研究于2011年3月-2012年2月对普定喀斯特区6个演替阶段(稀灌草丛、藤刺灌丛、灌木林、乔灌过渡林、次生乔木林以及原生乔木林)植物群落的凋落物生物量、组分(叶、枝、花果等)、月动态进行了定位观测.结果表明:6个演替阶段群落年均总凋落量分别为789.5、3821.8、4315.0、4158.8、4201.0和3950.4 kg·hm-2;随着群落正向演替的推进,植物群落凋落物量表现为先升高后缓慢下降最后趋于稳定.叶凋落量占总凋落量的59.0％ ～ 77.7％,枝凋落量占7.9％～24.0％,花果等凋落量占5.4％ ～ 24.0％,叶凋落量比例随群落正向演替递减,枝凋落量与群落演替阶段显著正相关.原生乔木林、灌木林和稀灌草丛群落凋落物的月变化规律为单峰型,其余3个演替群落凋落物的月变化规律均为双峰型;叶凋落物的月动态与总凋落物一致.     

黔西北地区不同演替阶段植物群落结构与物种多样性特征

该文采用"空间代替时间"的方法,研究了贵州省威宁县喀斯特地区植被演替过程中的群落结构、物种组成、生活型谱和物种多样性的变化规律。结果表明:(1)该调查共记录到种子植物174种,隶属于52科117属,物种分布较多的有菊科、蔷薇科、禾本科、杜鹃花科、小檗科、唇形科、蓼科。(2)随着植被的正向演替,物种丰富度逐渐增加,群落结构趋于复杂,高位芽植物所占比例逐渐增大。(3)随着植被的恢复,群落层次分化逐渐明显,大径级植株所占比例呈现增加趋势。(4)随着植被的恢复,群落各层次的ShannonWiener多样性指数(H)、Simpson多样性指数(DS)、均匀度指数(J)和Margalef丰富度指数(DM)逐渐增加;不同演替阶段植物群落之间的Srensen相似系数呈现先上升后下降的趋势,Cody指数则表现为逐渐增加的趋势。黔西北地区不同演替阶段植物群落结构和物种多样性不同,建群种和关键种发生了明显变化,不同演替阶段植物群落结构和物种多样性的研究对喀斯特地区植被演替规律的认识和生态恢复具有重要意义。     

深圳湾福田红树林群落演替

对深圳湾福田凤塘河红树林演替过程中的群落结构、物种组成、多样性变化及演替趋势的研究结果表明:(1)演替各阶段群落均由秋茄(Kandelia candel)、木榄(Bruguiera gymnorrhiza)、桐花树(Aegiceras corniculatum)、白骨壤(Avicennia marina)及老鼠簕(Acanthus ilicifolius)5个种类组成。(2)演替早期(4 a)群落高度增长较快,之后继续保持较快增长,至17 a时趋于稳定,之后增长速度缓慢。(3)4 a和17 a的群落立木级均分布在Ⅱ级—Ⅳ级,56 a和73 a的群落立木均达到最高级Ⅴ级,该两群落立木在Ⅱ级—Ⅴ级均有分布。(4)随着演替的进展,除了73 a群落盖度较56 a群落稍有下降外,其它演替群落盖度逐渐增加;群落胸高断面积表现出与盖度一致的变化趋势;群落个体密度则逐渐下降;群落物种多样性和均匀度不断提高,群落优势度则逐渐下降;群落相似性系数的变化表明在演替过程中群落物种组成结构没有发生明显变化。(5)群落变化及演替为:秋茄+桐花树群落→秋茄群落→白骨壤+秋茄群落→白骨壤群落。     

森林次生演替和土壤层次对微生物群落结构的影响

森林次生演替与生态系统结构和功能的动态变化密切相关。大多数研究主要关注植物群落以及土壤有机碳(SOC)的变化,然而土壤微生物群落如何响应森林次生演替还需要进一步探究。本研究以长白山森林次生演替序列(20、80、120、200和≥300年)和两个土壤层次为对象,采用磷脂脂肪酸微生物标志物,探究温带森林次生演替过程中地下微生物群落结构变化。森林次生演替改变了土壤微生物群落结构,主要归因于某些特定微生物类群的变化,演替前期革兰氏阴性菌和腐生真菌占主导,而在演替后期革兰氏阳性菌和丛枝菌根真菌占主导。另外,土壤有机质数量和质量差异是影响微生物群落结构和生物量的主要环境因素。森林演替前期和中期增加的SOC含量促进了微生物生物量,而演替后期增加的难分解芳香族有机组分抑制了微生物生物量合成。土壤层次间理化性质的差异导致微生物群落变化,有机质层高的SOC以及氮含量导致更多微生物生物量的合成。微生物群落在时间和空间尺度的变化及其驱动因素反映了生态系统结构和功能对环境变化的响应。     

群落结构和叶面积指数在具茨山植被次生演替中的变化

采取3S(GIS,RS和GPS)技术和野外作业相结合的方法,研究了河南省具茨山人为干扰后植被次生演替过程。在此基础上,从宏观和微观尺度对不同演替阶段典型群落的结构变化和植被叶面积指数(LAI)动态进行分析。结果表明:乔木阶段的植被LAI均值为4.1～5.5;灌木阶段为3.0～3.7;草本阶段为1.0～1.5。部分灌木林在生长季的LAI和盖度高于乔木林。随着演替进行,群落结构也发生很大变化,乔木种类和数量逐步上升。植被系统的复杂度和稳定性不断增强,其发挥的生态功能和生态服务价值随之提高。     

森林次生演替和土壤层次对微生物群落结构的影响

森林次生演替与生态系统结构和功能的动态变化密切相关。大多数研究主要关注植物群落以及土壤有机碳(SOC)的变化,然而土壤微生物群落如何响应森林次生演替还需要进一步探究。本研究以长白山森林次生演替序列(20、80、120、200和≥300年)和两个土壤层次为对象,采用磷脂脂肪酸微生物标志物,探究温带森林次生演替过程中地下微生物群落结构变化。森林次生演替改变了土壤微生物群落结构,主要归因于某些特定微生物类群的变化,演替前期革兰氏阴性菌和腐生真菌占主导,而在演替后期革兰氏阳性菌和丛枝菌根真菌占主导。另外,土壤有机质数量和质量差异是影响微生物群落结构和生物量的主要环境因素。森林演替前期和中期增加的SOC含量促进了微生物生物量,而演替后期增加的难分解芳香族有机组分抑制了微生物生物量合成。土壤层次间理化性质的差异导致微生物群落变化,有机质层高的SOC以及氮含量导致更多微生物生物量的合成。微生物群落在时间和空间尺度的变化及其驱动因素反映了生态系统结构和功能对环境变化的响应。     

普洱季风常绿阔叶林次生演替中木本植物幼苗更新特征

以时空替代的方法,将针阔混交林、季风常绿阔叶林的次生林与成熟林等3个处于同一空间下的群落作为次生演替进程中的3个阶段,研究云南普洱地区次生演替过程中的木本植物幼苗更新特征,分析了次生演替过程中木本植物幼苗的物种组成、密度、高度级及与环境因子的相关性。结果表明:在8个共144 m2的幼苗样地中调查木本植物幼苗101种2014株,其中乔木幼苗是主要组成。随着次生演替的进行,木本植物幼苗、乔木与藤本幼苗密度逐渐增加,灌木幼苗密度无显著变化;藤本植物幼苗的物种丰富度随着次生演替进行而增加,乔木与灌木幼苗则无显著变化,成熟季风常绿阔叶林中木本植物幼苗Shannon-Wiener指数要显著小于针阔混交林与次生季风常绿阔叶林。次生与成熟季风常绿阔叶林木本植物幼苗多度随高度级增加而减少,针阔混交林则呈现偏峰曲线,幼苗密度均集中分布在高度20 cm以内,3个群落演替阶段木本植物幼苗物种丰富度随高度级增加呈现偏锋曲线。相似性系数反映出乔木和藤本幼苗的更新来源与群落的物种组成存在着紧密的联系。乔木幼苗密度分布与样地坡度之间存在着显著的负相关,灌木幼苗密度与土壤pH值之间存在着显著正相关。     

云南松林次生演替阶段土壤细菌群落的变化

土壤细菌多样性是维持森林生态系统功能的关键因子,森林演替是影响其动态变化的重要因素。研究云南松林不同演替阶段土壤细菌群落结构及其多样性的变化规律,有助于深入理解森林生态系统恢复过程的驱动机制。本研究以云南省永仁县皆伐后形成的针叶林、针阔混交林和常绿阔叶林为对象,基于Illumina Hiseq高通量测序技术,分析森林演替过程中土壤细菌群落组成、结构、多样性及其影响因子的变化。结果表明: 土壤细菌的种群分类单元、Ace指数、Chao1指数和Shannon指数均随着演替进行呈减少趋势,演替早期阶段土壤的细菌总数、菌群丰富度及复杂程度最高。不同演替阶段细菌群落结构存在显著差异,其中,针阔混交林的差异最大,变形菌门和酸杆菌门为各演替序列共有的优势类群,放线菌门、绿弯菌门和Patescibacteria是演替早期的优势类群,且随着演替进行呈现减少趋势;变形菌门和WPS-2相对多度随演替进行呈增加趋势。土壤pH和乔木层物种丰富度是驱动次生演替过程中土壤细菌群落组成变化的关键因子。随着演替的进行,土壤细菌多样性减少,群落组成差异加大。     

The Community Dynamics of the Forest Secondary Succession in Heishiding Natural Reserve of Guangdong Province

Studying on the community structure, species composition and species diversities of the secondary succession of lower subtropical forest in Heishiding Natural Reserve, Guangdong Province, the following results were obtained. In 2 ~4 years of restored stage the seedlings of pioneer species, heliophytes and mesophytes occur simultaneously. In 10 years of restored stage pioneer species are superior as heliophytes maintain stable and mesophytes are being decreased. In the stage of mixed forest pioneer species are on the decreasing and heliopytes are on the increasing trend. In the stage of evergreen broad-leaved forest dominated by heliophytes pioneer species are being declined, heliophytes are dominant and mesophytes are being increased. In the stage of evergTeen broad-leaved forest dominated by mesophytes heliophytes are being declined and mesophytes are superior. The vertical structure, horizontal structure, the structure of species composition determine stability of communities in secondary succession process. The vertical structure of community is building up in the stage of restoration. The community is composed of the stands under the third class. The forest canopy is not stratified. The individual density is stable. From restored stage to mixed forest the vertical structure of community is diversified. There are the third and forth class stands in community. The tree layer is divided into three sub-layers. The individual density declines rapidly because of self-thinning. In the mixed forest the vertical stmcture is relatively stable. The abundance of the fifth class stands and area of breast height reach the first peak in the succession serial. The individual density increases rapidly. In stage of evergreen broad-leaved forest dominated by heliophytes, the vertical structure changes largely. The abundance of the fifth class and area of breast height decline rapidly, but the individual density changes slowly. In the stage of evergreen broad-leaved forest dominated by mesophytes, the vertical structure of community is stable and the composition of every stand class becomes more stable. The area of breast height is the biggest in the succession serial. When succession develops, the coverage of communities increases gradually but the turnover rate of species declines. The species composition tends to be stabilized. The diversity of species and community evenness increase gradually thus, the ecological dominance declines gradually.     

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.     

树种组成和径级结构对热带次生林生物量恢复影响的研究

本研究在海南尖峰岭和吊罗山热带林区海拔245～1255 m范围内根据林分恢复时间设立固定监测样地,探讨了森林地上生物量与树种组成和径级结构的关系.结果显示:海南热带次生林平均地上生物量为(155.38±37.16)×103 kg/hm2,其中低地次生雨林为(137.91±31.02)×103 kg/hm2,山地次生雨林...     

黄土高原植被不同演替阶段优势种的光合生理特性

在黄土高原子午岭林区,按照植被的演咎序列,分别选择植被次生演替不同阶段的优势种-白羊草、铁杆蒿、沙棘、狼牙刺、山杨和辽东栎,对其土壤与叶片氮素含量、叶片气体交换参数、叶绿素含量及叶绿素荧光参数进行了观测．结果表明：随着植被演替,0—20cm土壤全氮含量呈增大趋势,植被优势种叶片氮含量先升高后降低,叶绿素含量变化与优势种叶片氮含量相似．灌木群落优势种（沙棘、狼牙刺）叶片氮含量明显高于其它群落优势种（P〈0．05）,草本群落优势种（白羊草、铁杆蒿）和灌木群落优势种的光合速率（Pn）高于早期森林群落优势种（山杨）和顶级群落优势种（辽东栎）,叶片气孔导度（ga）与Pn变化规律相似．草本群落和灌木群落优势种叶片蒸腾速率（Tr）较高,随着演替的进行t逐步下降,顶级群落优势种辽东栎Tr较低．PSⅡ最大光化学量子效率（Fv／Fm）值呈缓慢增长趋势,辽东栎〉山杨〉沙棘〉狼牙刺〉铁杆蒿〉白羊草,而PSⅡ量子效率（ΦPSⅡ）值呈先增加后降低趋势;光化学淬灭系数（qp）在演替过程中呈整体增加趋势,而非光化学淬灭系数（qNP）呈先增加后降低趋势．这说明植被不同演替阶段优势种的光合生理特征以及对微生境的适应性存在明显差异．     

Structural changes of vegetation and its association with microclimate in a successional gradient of low thorn forest in northeastern Mexico

Understanding how microclimate and vegetation are associated during secondary succession is of primary importance for plant conservation in the face of the increasing land cover modification. However, these patterns are still unstudied for many plant communities. This study aimed to evaluate the structure (species richness, Shannon's diversity index, Simpson´s dominance index, abundance of each species, average height of species, species cover (%), species composition, and indicator values) of a low thorn forest fragment and to analyze its relation with microclimate along a successional gradient. Four stages of succession were delimited by the analysis of Landsat images, in the state of Tamaulipas, northeast Mexico. Statistical models incorporated species richness, diversity indices, abundance, height, and cover, as variables for searching differences between stages, or to evaluate microclimate associations. A total of 70 species, 54 genera, and 27 families were determined. Height of tree layer was the most important variable for discrimination of the successional stages. Conserved areas differed floristically from other stages, associated mainly with the lowest values of wind speed originated by tree layer characteristics. A significant association between species and microclimate was found, being wind speed and relative humidity the most important variables. Some species, due to their high importance values and their patterns of association with microclimate, may be considered as key taxa for low thorn forest, which is a threatened semitropical community in northeast Mexico. Conserved and late successional areas account for climatic regulation of this plant community, and the importance of these forest patches may be considered when establishing biodiversity protection areas.

     

岷江冷杉林皆伐后次生群落结构和物种多样性的演替动态

为阐明岷江冷杉林皆伐后次生群落的演替过程,采用空间代替时间的方法,在川西米亚罗林区海拔3100—3600 m的阴坡选择岷江冷杉林皆伐后次生演替10、20、30、40和50a阶段的次生群落作为研究对象,对其群落结构和物种多样性的动态进行了研究。不同演替阶段的树木均呈显著聚集分布。按群落中优势种的重要值将该演替序列划分为3个类型:悬钩子-蔷薇灌丛、白桦阔叶林和桦木-岷江冷杉针阔混交林。随次生演替,乔木和灌木种的物种丰富度趋于增加,而草本种的物种丰富度趋于减少;乔木和灌木种的Shannon-Wiener多样性指数趋于增大,而草本的Shannon-Wiener多样性指数趋于减小;乔木、灌木和草本层的Pielou均匀度指数均趋于增大;乔木层的Simpson优势度指数趋于减小,灌木和草本层的Simpson优势度指数在演替0—40年阶段趋于增大,而在演替50a阶段趋于减小。在该演替序列中,乔木、灌木和草本层的物种组成均呈耐荫种替代非耐荫种的趋势。     

Plant–plant interactions change during succession on nurse logs in a northern temperate rainforest

Plant–plant interactions change through succession from facilitative to competitive. At early stages of succession, early‐colonizing plants can increase the survival and reproductive output of other plants by ameliorating disturbance and stressful conditions. At later stages of succession, plant interactions are more competitive as plants put more energy toward growth and reproduction. In northern temperate rainforests, gap dynamics result in tree falls that facilitate tree regeneration (nurse logs) and bryophyte succession. How bryophyte‐tree seedling interactions vary through log succession remains unclear. We examined the relationships of tree seedlings, bryophyte community composition, bryophyte depth, and percent canopy cover in 166 1.0 m² plots on nurse logs and the forest floor in the Hoh rainforest in Washington, USA, to test the hypothesis that bryophyte‐tree seedling interactions change from facilitative to competitive as the log decays. Tree seedling density was highest on young logs with early‐colonizing bryophyte species (e.g., Rhizomnium glabrescens) and lowest on decayed logs with Hylocomium splendens, a long‐lived moss that reaches depths >20 cm. As a result, bryophyte depth increased with nurse log decay and was negatively associated with tree seedling density. Tree seedling density was 4.6× higher on nurse logs than on the forest floor, which was likely due to competitive exclusion by forest floor plants, such as H. splendens. Nurse logs had 17 species of bryophytes while the forest floor had six, indicating that nurse logs contribute to maintaining bryophyte diversity. Nurse logs enable both tree seedlings and smaller bryophyte species to avoid competition with forest floor plants, including the dominant bryophyte, H. splendens. H. splendens is likely a widespread driver of plant community structure given its dominance in northern temperate forests. Our findings indicate that plant–plant interactions shift with succession on nurse logs from facilitative to competitive and, thus, influence forest community structure and dynamics.     

Temporal variation in the influence of forest succession on caterpillar communities: A long‐term study in a tropical dry forest

Forest succession can influence herbivore communities through changes in host availability, plant quality, microclimate, canopy structure complexity and predator abundance. It is not well known, however, if such influence is constant across years. Caterpillars have been reported to be particularly susceptible to changes in plant community composition across forest succession, as most species are specialists and rely on the presence of their hosts. Nevertheless, in the case of tropical dry forests, plant species have less defined successional boundaries than tropical wet forests, and hence herbivore communities should be able to persist across different successional stages. To test this prediction, caterpillar communities were surveyed during eight consecutive years in a tropical dry forest in four replicated successional stages in Chamela, Jalisco and Mexico. Lepidopteran species richness and diversity were equivalent in mature forests and early successional stages, but a distinctive caterpillar community was found for the recently abandoned pastures. Species composition tended to converge among all four successional stages during the span of eight years. Overall, our results highlight the importance of both primary and secondary forest for the conservation of caterpillar biodiversity at a landscape level. We also highlight the relevance of long‐term studies when assessing the influence of forest succession to account for across year variation in species interactions and climatic factors. Abstract in French is available with online material.     

Pathways, mechanisms and predictability of vegetation change during tropical dry forest succession

The development of forest succession theory has been based on studies in temperate and tropical wet forests. As rates and pathways of succession vary with the environment, advances in successional theory and study approaches are challenged by controversies derived from such variation and by the scarcity of studies in other ecosystems. During five years, we studied development pathways and dynamics in a chronosequence spanning from very early to late successional stages (ca. 1–60 years) in a tropical dry forest of Mexico. We (1) contrasted dynamic pathways of change in structure, diversity, and species composition with static, chronosequence-based trends, (2) examined how structure and successional dynamics of guilds of trees shape community change, and (3) assessed the predictability of succession in this system. Forest diversity and structure increased with time but tree density stabilized early in succession. Dynamic pathways matched chronosequence trends. Succession consisted of two tree-dominated phases characterized by the development and dynamics of a pioneer and a mature forest species guild, respectively. Pioneer species dominated early recruitment (until ca. 10 years after abandonment), and declined before slower growing mature-forest species became dominant or reached maximum development rates (after 40–45 years). Pioneers promoted their replacement early in succession, while mature-forest species recruited and grew constantly throughout the process, with their lowest mortality coinciding with the peak of pioneer abundance. In contrast to prevailing stochastic views, we observed an orderly, community driven series of changes in this dry forest secondary succession. Chronosequences thus represent a valuable approach for revealing system-specific successional pathways, formulating hypotheses on causes and mechanisms and, in combination with repeated sampling, evaluating the effects of vegetation dynamics in pathway variation.