林窗与生物多样性维持

自 2 0世纪以来 ,森林群落演替更新与生态系统的空间异质性 (ｓｐａｔｉａｌｈｅｔｅｒｏｇｅｎｅｉｔｙ) ,逐渐成为当前生态学研究的热点。干扰 (自然或人为干扰 )作为群落时空格局及其动态的驱动力 ,是森林群落发展变化和结构维持的必要成分。将干扰状况与森林群落特征结合起来的研究方法 ,有助于对群落动态变化规律的深入理解 ,从而加深人们对森林群落调控机理的全面认识。生物多样性是全球关注的热点问题之一。对于生物多样性的维持与保护 ,提出了众多的理论与假说 ,但尚未形成一个完善的理论体系。空间异质性、物种竞争、边缘效应等都…     

红松老龄林主要树种的空间异质性特征与比较的定量研究

 空间异质性研究对认识不同尺度上的生态系统结构、功能和过程具有重要的理论意义。采用地统计学理论和方法,通过空间异质性特征和比较研究了我国东北红松老龄林中主要树种,即红松(Pinus koraiensis)、云杉(Picea koraiensis)、冷杉(Abies nephrolepis)、枫桦(Betula costata)和紫椴(Tilia amurensis)的空间异质性程度、空间异质性组成、尺度和格局问题。结果表明：红松老龄林中主要树种虽然生长在同一林分中,但是它们的空间异质性与格局具有明显的差异。红松的空间异质性程度最高,分数维较低,其空间异质性主要由空间自相关部分组成,在450m的尺度内,具有明显的空间格局。枫桦的空间异质性程度最低,分数维最大,空间异质性主要由随机部分组成,在大于100m的尺度上不存在明显的空间格局。云杉和冷杉由于生长的立地相似,因此具有相似的空间异质性特征和格局。紫椴是红松最重要的伴生树种,其空间异质性程度、空间异质性组成、尺度和格局等方面与红松接近。红松老龄林中主要树种空间异质性与格局在一定尺度上的差异,反映这些树种在生态系统中具有不同的生态学功能和过程。     

不同采伐强度对阔叶红松林主要树种空间分布格局和物种空间关联性的影响

分析采伐后森林群落中物种的空间格局有助于认识该格局形成的生态学过程、种群的生物学特性及其与环境因子之间的相互关系,并对制定可持续的森林经营方案具有重要意义。以长白山地区经历不同采伐方式形成的阔叶红松林次生林为研究对象,利用空间点格局分析的研究方法,探讨了采伐对阔叶红松林主要树种空间分布格局、种间关联性以及更新的影响。研究结果显示:较低强度的择伐对阔叶红松林主要树种的空间分布格局、种间关联性的改变较小,群落可以在较短时间内恢复。中等强度的择伐减少了成年树种对幼树的抑制作用,可以促进森林的天然更新。皆伐后,森林的群落结构,物种的空间分布格局、种间关联性都发生显著变化,尽管更新状况良好,但要恢复到伐前水平仍需要较长时间。择伐不仅通过改变主要树种的密度对阔叶红松林群落结构产生影响,还通过改变物种空间关联性改变群落的结构动态。因此,在制定森林生态系统经营管理方案时,不仅要选择适合的采伐强度,还要综合考虑采伐时物种的选择以及种间关系。     

气候变化对森林演替的影响

森林演替是森林生态动力源驱动下森林再生的生态学过程,自20世纪初建立群落演替理论以来,演替研究成为生态学研究中的热点．客观准确地认识森林演替规律,研究森林演替动力学机理及其模型,是科学管理森林生态系统的需要;对于天然林保护工程与森林植被的恢复重建,具有重要的理论与实际意义．干扰是森林循环的驱动力,导致森林生态系统时空异质性,是更新格局和生态学过程的主要影响因素．它可改变资源的有效性,干扰导致的林隙是森林循环的起点．回顾了目前演替研究的几种方法,即马尔科夫模型、林窗模型(GAP)、陆地生物圈模型(BIOME)和非线性演替模式．介绍了气候变化对森林演替的影响;并在已有成果的基础上,提出了目前研究存在的问题及未来的发展方向．     

辽东山区次生林木本植物空间分布

森林木本植物的空间格局有助于揭示群落结构的形成机制与潜在的生态学过程,且对林分经营具有一定指导意义。在0—50 m尺度范围内综合分析了辽东山区4 hm2温带次生林样地多度10的树种空间格局。研究发现:(1)在完全随机零模型下,大部分树种呈现聚集格局,聚集格局树种的比例随尺度增加而降低;在32 m的较大尺度下,随尺度增加,随机和规则格局成为树种分布的主要形式;(2)在异质性泊松过程零模型下,55.9%的树种呈现随机格局,其余大部分树种在10 m的尺度下呈现聚集格局,且随尺度增加,规则格局成为主要形式;(3)在完全随机零模型下,树种属性(林层、径级和多度)显著地影响种群聚集度,而在异质性泊松过程零模型下,树种属性对种群聚集度不存在显著影响。综上,生境异质性、扩散限制和树种属性部分解释了辽东山区次生林木本植物空间分布格局,相对而言,生境异质性的效应更为突出。研究结果有助于揭示次生林群落生物多样性的维持机制。     

空间和平衡理论在景观生态学中的发展和应用

传统生态学承认生态系统中平衡、稳定状态的存在 ,“自然平衡理论”是其集中体现。而近 30年来 ,非平衡理论的出现 ,使得人们对生态系统的认识发生了彻底的改变。非平衡理论认为 ,偶然事件 (如干扰等 )在生态过程中起着重要作用。随着认识的转变 ,对生态过程中的“空间”和空间异质性的理解也发生了变化 ,传统生态学总是试图回避空间动态和空间异质性问题 ,而着重强调时间格局的重要性。但在近 2 0年来 ,空间格局则得到了广泛重视。空间理论及空间格局的研究表明 ,空间异质性既可能增加生态系统和生态过程的稳定性 ,也可能降低其稳定性。分析两种变化趋势 ,指出了它们的相互关系 ,同时讨论了空间理论的发展及其在景观生态学中的应用 ,并阐述了生态平衡的含义。     

林下密集蕨类层生态学研究进展

林下密集蕨类层在森林尤其是受干扰森林中广泛存在,对森林更新具有强烈的过滤效应,能够改变林下层的多样性,影响群落的结构、功能和动态,林下密集蕨类层的生态学研究,对揭示森林群落物种分布格局和群落构建机制具有重要意义。结合国内外最新研究,论述了林下密集蕨类层的特征,从机制上解释了林下密集蕨类层形成的主要原因;分类阐述了林下密集蕨类层通过资源竞争、化学互感、动物取食策略和机械干扰等途径影响森林更新的过程;探讨了林下密集蕨类层对森林演替动态的潜在影响。最后,简要指出了目前林下密集蕨类层研究中存在的一些不足。     

景观生态学发展现状

景观生态学主要研究空间格局和生态过程的相互作用,与生态学有着良好的结合。过去的15年,景观生态学得到迅猛发展。土地利用历史和自然干扰对目前生态系统具有持久影响。格局指标（pattern metrics）的发展已保持相对稳定,并且广泛应用于反映景观格局与生态响应的关系。在研究生物类群的景观格局时,多尺度分析具有重要意义,同时,物种间的空间联系也日益受到生态学家的关注。干扰是景观生态学研究的一个重点,当前研究注重干扰的相互作用。尽管景观生态学与系统生态学之间的结合仍然面临挑战,但对景观功能的认识仍需要加强。景观生态学应继续立足于空间异质性进行研究,严格验证有关概念的通用性,同时更加深刻地认识景观格局与过程关系的机理     

林分光照空间异质性对水曲柳更新的影响

 光照是森林更新动态中极为关键的环境因子。本研究在东北天然次生林中选择样地,应用地统计学理论结合实验生态学方法,定量分析林分光照空间异质性特征,同时进行林下播种水曲柳（Fraxinus mandshurica）的更新实验,探测更新格局特征。通过空间关联性分析,研究探讨是否林分光照的空间变异对水曲柳更新有重要影响作用。研究结果表明： 1）生长季林分平均光照仅为全光照的4.2%～4.6%,在空间分布上呈现非常明显的异质性特征,空间自相关变异（62.5%～78.2%）主要体现在10.9～12.4 m的尺度上。2）林分光照不同的空间样点上,水曲柳更新幼苗发生数量和秋季幼苗存活数量均存在显著差异,光照相对强的微生境中,更新苗发生数量较多且存活率高,而在光照较弱的微生境中则更新较差,更新具有十分明显的空间格局。3）更新格局与林分光照空间异质性特征紧密关联。在光照空间异质性程度较高、空间变异较复杂的条件下,更新相应呈现出较复杂的空间格局,自相关的变异表现在较小尺度范围（2.12～6.97 m）。在光照空间异质性程度较低、空间变异复杂性较小的条件下,更新格局的复杂程度明显变小,空间自相关变异（>83%）表现在较大的尺度上（30 m）,对更新格局起主要的影响作用,随机变异的影响很小（<17%）。这些研究结果说明,林分光照环境在水曲柳更新中的作用十分关键,林分光照的空间异质性对更新格局起决定性作用。     

黑龙江省大兴安岭林区火烧迹地森林更新及其影响因子

林火干扰是大兴安岭森林更新的影响因子之一,研究火烧迹地森林更新的影响因子(立地条件、火前植被、火干扰特征)对理解生态系统的结构、功能和火后演替轨迹具有重要意义。选取呼中及新林林业局55个代表性火烧样地,利用增强回归树分析法分析了火烧迹地森林更新的影响因素。结果表明:(1)立地条件是影响针、阔叶树更新苗密度的主要因素;海拔对针叶树更新苗密度的影响最大;坡度对阔叶树更新苗密度影响最大;(2)距上次火烧时间对针叶树更新苗比重影响最大,其次是林型;(3)中度林火干扰后森林更新状况好于轻度和重度火烧迹地。根据火烧迹地森林更新调查分析可知:林型影响火后演替模式,火前为针叶树或阔叶树纯林,火后易发生自我更新(火后树种更新组成与火前林型相同),而针阔混交林在火干扰影响下易于发生序列演替(火后初期以早期演替树种更新为主)。     

长白山次生杨桦林种群空间点格局及密度制约效应

以长白山5.2 hm²次生杨桦林样地为研究对象,利用空间点格局分析的双相关函数g（r）以及随机标签零模型和案例-对照设计法,探讨了树木种群空间格局及其密度制约效应。研究结果表明：生境异质性对不同生活型以及不同树种的成熟个体在大尺度范围上具有强烈影响。剔除生境异质性带来的影响后,在14个常见树种中有12个树种在小尺度上呈显著的空间聚集分布格局;随着空间尺度的增加聚集性分布树种数量急剧下降,在18 m尺度上聚集率下降到0。全部14个常见树种均呈现出显著的密度制约效应,表明密度制约是调节温带森林树木种群空间结构的主要作用机制。此外,密度制约与物种多度呈负相关,与亚林层和灌木层树种的同种聚集强度呈正相关。并且随着空间尺度增大受密度制约影响的树种百分比逐渐减小,14个常见树种中有11个在0-1 m处达到最大的密度制约强度。     

辽东山区次生林乔木幼苗分布格局与种间空间关联性

在辽东山区次生林建立4 hm^2样地(200 m×200 m),研究0~50 m尺度范围内乔木幼苗分布格局及种间空间关联性.结果表明:在完全随机零模型下,0~20 m尺度上,95%的树种呈现聚集分布格局;0~16 m尺度上,19个树种呈现聚集分布;随着尺度的增加,聚集分布树种的比例逐渐减少,50 m尺度上,随机分布成为树种分布的主要形式;在异质性泊松过程零模型下,0~24 m尺度上,5%的树种呈现聚集分布,26~50 m尺度上,42%和58%的树种呈现随机和均匀分布.在完全随机零模型下,正相关树种对比例最高,且在50 m尺度下呈现正相关、负相关、无相关3种相关性的树种对比例相同;在异质性泊松过程零模型下,树种对主要呈现负相关,且随尺度增大,负相关的树种对比例逐渐升高.种子扩散限制和生境异质性在某种程度上解释了乔木幼苗的聚集分布格局,乔木幼苗强烈的聚集分布又促使种间空间关联性密切,更新群落稳定性较差.     

Spatial analysis of structural and tree‐ring related parameters in a timberline forest in the Italian Alps

Abstract. We investigated the variability in spatial pattern of some structural, dendrochronological and dendroclimatological features of a mixed Larix decidua‐Pinus cembra forest at the timberline in the eastern Italian Alps at fine geographical and temporal scales. Forest structure variables such as stem diameter, tree height, age and tree‐ring related parameters (yearly growth index, mean sensitivity, first order autocorrelation and some dendroclimatic variables) have been compared at various scale levels. We observed that most of the variables show positive autocorrelated structures due to both forest dynamics and fine‐scale driving forces, probably related to microrelief. Spatial structure of yearly indexed radial growth appears sensitive to extreme climatic events. Secondary succession after past disturbances drives the forest towards a structure governed by a gap regeneration dynamics that seems to ensure the different requirements of the two main tree species present. Small spatial scale studies of forest structures, especially if integrated to dendro‐ecological data, seem an efficient tool to assess the disturbance regime and species sensitivity to environmental change.     

Soil heterogeneity in tree mixtures depends on spatial clustering of tree species

Heterogeneity in soil characteristics promotes and maintains coexistence between a diverse set of species. In forests, trees have species-specific impacts on soil abiotic characteristics and mixing of tree species is being promoted as a tool to ensure high levels of diversity and functioning. Yet, limited knowledge is available on the effect of tree species composition and spatial clustering on heterogeneity in soil characteristics. In this paper we derived heterogeneity of key characteristics of the leaf litterfall, the forest floor and the mineral topsoil (C, N and base cation concentration, C:N ratio and mass) in 53 plots of 7 different tree species compositions. We found that heterogeneity increased from the leaf litterfall, through the forest floor down to the mineral topsoil. Mixing tree species did not lead to an increased heterogeneity in the forest floor and topsoil compared to monocultures. However, we did find that mixed plots where conspecific trees stand in groups are more heterogeneous than plots where species are intimately mixed. Our results imply that heterogeneity in soil characteristics does not necessarily increase with tree diversity, but that within mixed stands the spatial organization of tree species should be considered in relation to the scale at which heterogeneity is desired.     

大兴安岭次生林空间分布格局及其尺度效应

研究大兴安岭落叶松林、白桦林和落叶松+白桦混交林3种次生林中,不同树种(落叶松、白桦和其他)和不同大小级(Ⅰ～Ⅴ级)树木的空间分布格局及其尺度效应。结果表明:3个林型中,仅落叶松林的更新数量达到良好状态,其余2种林型均为更新不良;各林型中,更新层幼苗、幼树的数量组成均与乔木层存在显著差异,且各林型中树木的直径(落叶松林和落叶松+白桦混交林除外)和树高分布也不够合理,3个林型均属于不稳定群落。各林型内林木空间分布整体以聚集分布为主,但方差/均值比率、负二项指数、Green指数、平均拥挤度和Morisita指数5个判别指数随尺度的变化显著不同,并以线性递增(40%)、幂函数递增(22%)和负二次多项式(20%)趋势为主。Ⅰ～Ⅲ级的更新层林木在不同尺度上以聚集分布为主,而乔木层(Ⅳ～Ⅴ级)在多种分布格局间波动,其对尺度的响应同样以线性递增(44%)、幂函数递增(15%)和负二次多项式(12%)为主。同一林型和取样尺度下,林木空间聚集程度整体随着林木大小级的增加呈线性下降趋势。各林型中,非优势树种的种群格局规模往往大于优势树种,而更新层林木空间格局规模大于乔木层。     

空间异质性对样地数据空间外推的影响

应用模型结合的方法模拟了3个空间异质性等级预案下反应变量(气候变化下景观水平的树种分布面积)的变化情况,并分析模拟结果在预案之间的差异性,探讨了环境空间异质性对样地观测到的树种对气候变化响应向更大空间尺度外推的影响.结果表明:空间异质性在一般情况下对样地数据向土地类型尺度外推没有影响,而对样地尺度外推到海拔带尺度的影响则有较复杂的情况.对于对气候变化不敏感的树种以及非地带性树种,空间异质性对样地数据向海拔带尺度外推没有影响;对于大多数对气候变化敏感的地带性树种而言,空间异质性对样地数据向海拔带尺度外推则有影响.     

Regeneration of Rhizophora mucronata (Lamk.) in degraded mangrove forest: Lessons from point pattern analyses of local tree interactions

Spatial structural patterns emerging from local tree interactions influence growth, mortality and regeneration processes in forest ecosystems, and decoding them enhance the understanding of ecological mechanisms affecting forest regeneration. Point-Patterns analysis was applied for the very first time to mangrove ecology to explore the spatial structure of Rhizophora mucronata regeneration in a disturbed mangrove forest; and the pattern of associations of juvenile–adult trees. R. mucronata trees were mapped in plots of 50 m × 10 m located at the seaward, central and landward edge along 50 m wide transect in the forest, and the mapped patterns were analysed with pair correlation and mark-connection functions. The population density of R. mucronata differed along the tidal gradient with the highest density in the central region, and the least near the shoreline. The study revealed that short distance propagule dispersal, resulting in the establishment of juveniles in closed distance to the mother trees, might not be the driving force for distribution of this species. The spatial structural pattern of R. mucronata population along tidal gradient showed a characteristic spatial aggregation at small scale, but randomly distributed as the distances become larger. There was a distinct spatial segregation between recruits and adult trees, and hence spatially independent. Though, adult–adult trees associations did not show a clear spatial segregation pattern; the recruit–recruit species associations exhibited significant clustering in space. Although habitat heterogeneity might be responsible for the local scale aggregation in this population, the effect of plant–plant conspecific interactions is more probable to inform the long-term structure and dynamics of the population of R. mucronata, and ditto for the entire forest.     

Structural heterogeneity and spatial autocorrelation in a natural mature Pinus sylvestris dominated forest

The tree species composition, vertical stratification and patterns of spatial autocorrelation at the tree and quadrate (25 × 25 m) scales were studied in a natural mature PinuS sylvestris dominated forest in eastern Finland. For the analyses we mapped the locations and dimensions of trees taller than 10 m in a 9 ha (300 × 300 m) area, and within this area we mapped all trees taller than 0.3 m on a core plot of 4 ha (200 × 200 m). The overall tree size distribution was bimodal. the dominant layer and the understory forming the peak frequencies. Pinus sylvestris dominated the main canopy, together with scattered Betula pendula and Picea abies. Alnus incana, Populus tremula, Salix caprea, Sorbus aucuparia and Juniperus communis occurred only in the under- and middlestories. Autocorrelation analysis (semivarianee) of tree size variation revealed spatial patterns, which were strongly dependent on the size of trees included in the analysis. When all living trees, including the understory regeneration, were taken into account, the autocorrelation pattern ranged up to 35 m inter-tree distances, reflecting the spatial scale of understory regeneration patches. Competitive interaction among middle- and upperstory trees (height>10 m) had contrasting effects on autocorrelation pattern depending on spatial scale. At the fine scale, dominant trees suppressed their smaller close neighbors (asymmetric competition), which was shown as increased tree size variation at small inter-tree distances (<2 m). At slightly larger inter-tree distances, specifically among large trees of similar size, competition was more symmetrical, which resulted in decreased tree size variation at these inter-tree distances (3–4 m). This effect was seen most clearly in the dominant trees, there being a clear autocorrelation pattern in tree size up to inter-tree distances of ～4 m. At the quadrate scale (25 × 25 m) the analysis revealed high local variation in structural characteristics such as tree height diversity (THD), tree species diversity (H) and autocorrelation of tree height. The analysis suggests that naturally developed P. sylvestris forests exhibit complex small-scale patterns of structural heterogeneity and spatial autocorrelation in tree size. These patterns may be important for stand-scale habitat diversity and can have aggregated effects on ecosystem dynamics at larger spatial scales though their influence on the spread of disturbance and regeneration after disturbance.     

Effects of heterogeneity of pre-fire forests and vegetation burn severity on short-term post-fire vegetation density and regeneration in Samcheok,Korea

This study investigated the combined effects of heterogeneity of pre-fire forest cover and vegetation burn severity on post-fire vegetation density and regeneration at an early stage in Samcheok, Korea. To measure the spatial heterogeneity of pre-fire forests, spatial pattern metrics at a landscape level and class level were adopted, and a regression tree analysis for post-fire vegetation density and regeneration was used to avoid spatial autocorrelation. Two regression tree models were estimated for post-fire vegetation density and post-fire vegetation regeneration with the same independent variable sets, including heterogeneity of pre-fire forest cover and vegetation burn severity. The estimated model suggested that the percentage of Japanese red pine and burn severity were the most significant variables for post-fire vegetation density and regeneration, respectively. The compositional and spatial heterogeneity of pre-fire forest and burn severity, as well as the degree of burn severity, was found to have significant impacts on post-fire vegetation density and regeneration. Overall, more rapid vegetation regeneration can be expected in more severely burned areas. However, this rapid vegetation regeneration at an early stage is due mostly to perennials and shrubs, not to the sprouting or regrowth of trees. The study results strongly indicated that a susceptible forest cover type and its spatial patterns directly influence the heterogeneity of burn severity and early vegetation density and regeneration. Hence, the management of susceptible forest cover types is particularly critical for establishing more fire-resilient forests and for post-fire forest restoration.