不同密度条件下芨芨草空间格局对环境胁迫的响应

基于小尺度上植物间相互作用与空间格局的高度相关性,选用可避免环境异质性影响的K2点格局函数,研究了沙枣-芨芨草群落中63个芨芨草样方(3密度条件×3生境条件×7重复)中芨芨草种群在小尺度下(0—0.5 m)的空间格局,探讨了在3种密度条件(高、中、低)及3种生境条件下(冠盖区、过渡区、空旷区),芨芨草种群空间格局对土壤理化性质胁迫(盐分、有机质及容重)程度的响应。研究结果表明,就土壤盐分,养分含量及土壤容重而言,土壤理化性质胁迫程度沿冠盖区、过渡区和空旷区增加的趋势。相应地,芨芨草种群呈聚集分布的样方比例在沿冠盖区(6/21)较低,而在过渡区(11/21)和空旷区(11/21)较高。然而,在不同密度条件下,芨芨草种群空间格局对土壤性质胁迫的响应不同。在低密度条件下,芨芨草种群在冠盖区多数为聚集分布(4/7),在过渡区和空旷区全部为聚集分布;中密度条件下,芨芨草聚集分布样方比例沿冠盖区-过渡区-空旷区梯度增加(分别为2/7,3/7,4/7),但低于同等胁迫条件下低密度样方中聚集分布数量;高密度条件下,在各胁迫条件下,除了一个过渡区高密度样方,其余芨芨草种群均为随机分布。总体上,随着环境胁迫增强,在中低密度下,芨芨草种群趋向于聚集分布;但在高密度下,芨芨草种群均以随机分布为主。此外,芨芨草种群空间格局随密度变化趋势比随土壤理化性质胁迫梯度变化趋势更加明显,可能表明相对于土壤理化性质胁迫程度,芨芨草种群密度对其空间格局影响更大。因此,在考虑芨芨草种群空间格局对环境胁迫的响应时,应当考虑种群密度因素。     

A spatial model of forest dynamics

Effects of spatial processes on temperate deciduous forest structure and dynamics were investigated with a spatial simulator derived from a forest gap model. The multi-species neighborhood model accounted for competitive interactions and endogenous disturbance in the form of small canopy gaps. Simulated and actual spatial pattern of old-growth stands were compared. The 400 yr simulations produced a pattern scale (0.07–0.2 ha patches) similar to that of an actual stand; simulated pattern intensity was greater than actual intensity, however. Distances to nearest neighbor were somewhat similar for trees in the simulated and actual stands; yet the frequency distributions of distance to nearest neighbor values differed substantially. The simulated stand patterns were generally less random than the actual patterns. Spatial pattern changed markedly during the course of simulated succession. Pattern approached a random dispersion in early succession. Intensity peaked at mid-succession (ca. 150 yr) with a hyperdispersed overstory and a strongly clumped understory. Pattern intensity diminished in late succession as a mixed size structure developed. Old-growth patch size was greater than the neighborhood (or gap) size, suggesting the gap-sized areas do not behave independently.     

植物空间分布格局中邻体距离的概率分布模型及参数估计

最近邻体法是一类有效的植物空间分布格局分析方法,邻体距离的概率分布模型用于描述邻体距离的统计特征,属于常用的最近邻体法之一。然而,聚集分布格局中邻体距离(个体到个体)的概率分布模型表达式复杂,参数估计的计算量大。根据该模型期望和方差的特性,提出了一种简化的参数估计方法,并利用遗传算法来实现参数优化,结果表明遗传算法可以有效地估计的该模型的两个参数。同时,利用该模型拟合了加拿大南温哥华岛3个寒温带树种的空间分布数据,结果显示:该概率分布模型可以很好地拟合美国花旗松(P.menziesii)和西部铁杉(T.heterophylla)的邻体距离分布,但由于西北红柏(T.plicata)存在高度聚集的团簇分布,拟合结果不理想;美国花旗松在样地中近似随机分布,空间聚集参数对空间尺度的依赖性不强,但西北红柏和西部铁杉空间聚集参数具有尺度依赖性,随邻体距离阶数增加而变大。最后,讨论了该模型以及参数估计方法的优势和限制。     

Seedling dynamics of <Emphasis Type="Italic">Acer mono</Emphasis> and <Emphasis Type="Italic">Fagus crenata</Emphasis>: an environmental filter limiting their adult distributions

Spatial pattern of a biotic population may change over time as its component individuals grow or die out, but whether this is the case for desert annual plants is largely unknown. Here we examined seasonal changes in spatial patterns of two annuals, Eriogonum abertianum and Haplopappus gracilis, in initial (winter) and final (summer) densities. The density was measured as the number of individuals from 384 permanent quadrats (each 0.5 m × 0.5 m) in the Chihuahuan Desert near Portal, Arizona, USA. We used three probability distributions (binomial, Poisson, and negative binomial or NB) that represent three basic spatial patterns (regular, random, and clumped) to fit the observed frequency distributions of densities of the two annuals. Both species showed clear clumped patterns as characterized by the NB and had similar inverse J-shaped frequency distribution curves in two density categories. Also, both species displayed a reduced degree of aggregation from winter to summer after the spring drought (massive die-off), as indicated by the increased k-parameter of the NB and decreased values of another NB parameter p, variance/mean ratio, Lloyd’s Index of Patchiness, and David and Moore’s Index of Clumping. Further, we hypothesized that while the NB (i.e., Poisson-logarithmic) well fits the distribution of individuals per quadrat, its components, the Poisson and logarithmic, may describe the distributions of clumps per quadrat and of individuals per clump, respectively. We thus obtained the means and variances for (1) individuals per quadrat, (2) clumps per quadrat, and (3) individuals per clump. The results showed that the decrease of the density from winter to summer for each plant resulted from the decrease of individuals per clump, rather than from the decrease of clumps per quadrat. The great similarities between the two annuals indicate that our observed temporal changes in spatial patterns may be common among desert annual plants.     

呼中林区火烧点格局分析及影响因素

林火是森林生态系统景观格局、动态和生态过程的重要自然驱动力,理解林火发生空间格局与影响因素对于林火安全管理具有重要的作用。采用点格局分析方法,以黑龙江大兴安岭呼中林区1990-2005年火烧数据为研究案例,分析了火烧点空间格局及其影响因素。结果表明,火烧点在空间上的分布是不均匀的,呈现聚集分布,存在一些火烧高发区和低发区。呼中林区火烧概率是0.004-0.012次/(km² · a),平均火烧概率为0.0077次/(km² · a)。人类活动因子、地形因子和植被因子对林火的发生均具有重要作用。应用空间点格局分析方法表明,距离居民点和道路的距离、高程、坡度和林型是影响林火发生的显著因子。因此在进行森林防火管理时,仅仅通过控制人类活动对于降低林火火险的效果是有限的,地形和林型也是林火防控时重点要考虑的因素。     

秦岭华山松种群分布格局及动态研究

基于相邻格子样方的野外取样方法,运用统计生态学的样方方差法、Ｇｒｅｉｇ－Ｓｍｉｔｈ格局分析法和Ｋｅｒｓｈａｗ格局强度分析法对秦岭林区不同年龄阶段华山松种群的空间分布格局、格局规模、格局强度进行了研究。     

山西翅果油树群落优势种群分布格局研究

 应用扩散系数、聚集指数、平均拥挤度、聚块性指数、Green指数、聚集强度、Poisson分布和负二项分布的X2拟合检验等方法,研究了山西翅果油树群落优势种群的分布格局,并用相关分析比较了6个指数间的关系,结果表明：翅果油树分布格局呈随机型,其余22个优势种的分布格局皆为聚集型,这主要与物种本身的生态和生物学特性有关,以及与物种的竞争排斥作用有密切联系。在判定物种分布格局的8种方法中,以方差／均值比率、Poisson分布和负二项分布的X2拟合检验联合运用效果较好,不仅生态学意义明确,而且结果具有严格的统计学意义。     

Neighbourhood effects influence drought‐induced mortality of savanna trees in Australia

Questions: The following hypotheses of neighbourhood effects on drought‐induced mortality are evaluated: (A) drought‐induced stem death is randomly distributed in space, (B) stems are predisposed to drought‐induced death through negative density‐dependent effects and (C) stems are predisposed to drought‐induced death due to local deficits in plant available resources. Location: Central Queensland, Australia. Methods: Recent mass mortality of woody stems was surveyed and mapped in three 1.21‐ha quadrats within Eucalyptus melanophloia‐dominated savanna. A multi‐faceted analytical approach was adopted including spatial pattern analyses, two logistic regressions of neighbourhood density effects on survival and spatial autocorrelation analyses of model residuals. Results: Mortality was concentrated in stems ≤15‐cm diameter at breast height (DBH). Survival was aggregated or random in quadrats 1 and 3 and random o regular in quadrat 2. Small neighbour density had a negative effect on survival in all quadrats. In addition, the second model identified a positive relationship between survival and living neighbour density in quadrat 3 (indicating a resource patch effect), but a negative relationship in quadrat 2 (density effect). Analysis of model residuals showed that neighbour density explained mortality equally well across quadrat 2, but not across quadrats 1 and 3. Conclusions: There was evidence in support of hypotheses B (neighbour density) and C (resource heterogeneity). We found strong support for an interaction between microsite quality and neighbourhood stem densities, and suggest that this interaction is driven by plant available water.     

Do spatial effects play a role in the spatial distribution of desert‐dwelling Acacia raddiana?

Abstract. We investigated the spatial pattern of A. raddiana in the Negev desert of Israel in order to gain insights into the factors and processes driving the dynamics of this species. Using a scale‐dependent measure, the ring statistic, we analysed both patterns observed in the field and time series of spatial tree distributions produced by a simulation model. In the field, random spacing was the predominant pattern observed. However seedlings were clumped on small scales. We ran the model under two contrasting scenarios representing hypotheses that explain the clumping of seedlings and the random distribution of trees. One hypothesis is that there is spatial heterogeneity in seed distribution, germination and seedling mortality, but that these heterogeneities are not correlated with each other in space. The second hypothesis assumes a correlation between these heterogeneities leading to areas suitable for establishment. However, the suitability of the sites is temporally variable. Furthermore, the second hypothesis assumes density‐dependent tree mortality due to competition. Both hypotheses lead to spatial distributions that are in qualitative agreement with the patterns observed in the field. Therefore, the classical view that a clumped seedling distribution and a random pattern of older trees is due to clumped regeneration and density‐dependent mortality may not hold for Acacia trees in the Negev.     

荒漠草原不同土壤条件下猪毛蒿幼苗种群的点格局分析

采用摄影定位法测定了宁夏荒漠草原3种不同土壤条件下猪毛蒿（Artemisia scoparia）幼苗种群的空间格局,并应用完全空间随机模型、泊松聚块模型和嵌套双聚块模型对其分布格局进行了分析。结果表明：（1）在灰钙土生境下,猪毛蒿幼苗种群在小尺度上（0-2.85m）表现为聚集分布,随着尺度的增大先呈现为随机分布（2.85-3.75 m）,然后又呈现为均匀分布（3.75-5m）;在风沙土生境下,猪毛蒿幼苗种群在0-1.85 m之间表现为聚集分布,在1.85-2.35 m之间表现为随机分布,当尺度大于2.35 m时表现为均匀分布;而基岩风化残积土上的猪毛蒿幼苗种群在整个尺度上均呈现随机分布。（2）猪毛蒿种群幼苗在基岩风化残积土上符合泊松聚块模型,即猪毛蒿种群空间格局的聚块中不存在较高密度的小聚块;而在风沙土和灰钙土上则符合嵌套双聚块模型,即在大聚块中分布较高密度的小聚块。猪毛蒿幼苗种群空间格局的形成与土壤异质性存在着密切的联系,种群在空间中分布格局的形成机制可以通过种群空间格局的分析加以解释。     

Population structure of Adenostoma fasciculatum in mature stands of chamise chaparral in the southern Sierra Nevada,California

Summary In the low elevation chaparral areas of Sequoia National Park, California, pure stands of chamise (Adenostoma fasciculatum) are periodically rejuvenated by fire. Mature stands showed considerable variability in density and total biomass even though a positive correlation exists between the two. Mature stands showed a preponderance of individuals in the smaller size classes (inverse-J shape distribution). Dead shrubs found in mature stands also tended to be in the smaller size classes. This relatively high mortality of small individuals is important to post-fire stand development. In addition, resprout and seedling biomass one year after fire both showed inverse-J shaped size-class structures. A positive correlation existed between the preburn basal area of a shrub and its first year resprout biomass. Shrub biomass and distance to nearest neighbor were poorly correlated. A significant correlation existed between stand density and a stand's variance-to-mean ratio, indicating a trend toward more regular spacing as density increases. Pre-burn and fire-induced mortality tended to move the stand towards a more clumped distribution. Seedlings replaced dead individuals after a fire and thus restored regular spacing.     

基于地理加权泊松模型的天然次生林进界株数空间分布与预测

基于帽儿山实验林场2004—2016年森林资源二类调查固定样地(共108块)数据,采用全局Poisson模型和4种空间尺度(2.5、5、10、15 km)下的地理加权泊松模型(geographically weighted Poisson regression, GWPR)对天然次生林进界株数的空间分布进行了研究,并对5种模型的拟合效果以及影响林分进界株数的因子进行了分析,利用莫兰指数描述了模型残差在全局和局域两种水平上的空间自相关性.结果表明: 本文所选的林分及地形因子都显著影响天然次生林进界株数的空间分布,林分平均胸径是最主要的影响因子;在小尺度(2.5 km)下GWPR模型拥有很高的拟合精度,产生了最大范围的模型参数估计值,得到了较好的模型参数局域化空间分布效果;在较小尺度(2.5和5 km)下GWPR模型产生了较小范围的模型残差,模型的稳定性得到提升;在小尺度(2.5 km)下GWPR模型残差的全局空间自相关性达到最低,局域空间自相关性显著减小,并形成了不同观测值少量聚类这一理想的空间分布模式;在对进界株数空间分布的模拟效果上,小尺度(2.5 km)下的局域模型明显好于全局模型.     

Structural segregation and scales of spatial dependency in Abies amabilis forests

Question: Is a mosaic structure apparent in the spatial distribution of trees in old‐growth Abies amabilis forests? Location: Montane forests of the western Cascade Range, Washington, USA. Methods: Maps of tree locations were created for study areas located in two, 300‐year old stands and a single 600‐year old stand. Stand structure parameters were calculated using several subsample quadrats sizes (56.25 ‐ 306.25 m²), which were drawn randomly with replacement at a density of 250 quadrats per ha from the stem maps in the computing environment. Spatial cross‐covariance functions between different canopy strata were estimated using the spline cross‐correlogram. Results: Negative spatial correlation (segregation) between subcanopy tree density and areas of high overstorey occupancy was detected. Understorey and midstorey tree densities were positively spatially correlated. These general trends were apparent across the range of observational scales investigated. Significant spatial correlation between canopy strata was observed at spatial scales of 12 ‐ 44 m and extended to the largest scales in the 600‐year old stand. Conclusion: The observed spatial segregation between canopy strata supports the hypothesis that old A. amabilis forests form fine‐scale structural mosaics. Structural segregation at small scales may be due to competitive interactions as well as exogenous forcing of tree locations (e.g. by mortality due to pathogens or disturbance), however segregation at large scales in the 600‐year old stand is likely due to exogenous factors alone. This study reinforces the idea that horizontal heterogeneity is an emergent property of old‐growth forests.     

广西木论保护区喀斯特常绿落叶阔叶混交林优势种空间格局及其相关性

基于广西木论国家级自然保护区喀斯特常绿落叶阔叶混交林1 hm²样地调查数据,采用点格局方法分析了混交林4个优势种群空间格局,及种内和种间的相关关系.结果表明：木论常绿落叶阔叶混交林乔木层Ⅰ亚层以小栾树为主,Ⅱ亚层和Ⅲ亚层均以东女贞、铁榄和广西密花树为主;4个优势种群不同林层的空间分布格局在<10 m尺度上呈显著的聚集分布,>25 m尺度呈随机分布,有些尺度呈均匀分布,10～25 m尺度内则为聚集分布向随机分布过渡,且不同树种聚集分布向随机分布转化的临界尺度不同.Ⅰ亚层小栾树与Ⅱ亚层4个优势种的相关性较弱;Ⅲ亚层各优势种与Ⅱ亚层4个优势种中,小栾树与4个优势种相关关系波动较大,东女贞<50 m尺度内呈显著正相关,铁榄与4个优势种没有明显的相关性,广西密花树在<20 m尺度没有相关性,而在20～50 m尺度上呈显著负相关.     

鄂尔多斯高原北缘水分梯度下天然植被分布格局初探

以鄂尔多斯高原杭锦旗北部独贵塔拉镇的过渡带为研究对象,通过样线调查,研究了水分梯度下天然植被的空间分布规律.TWINSPAN的分类结果表明,群落以中生和旱生为主,一些耐盐的植物散布其中;相邻样方间相似性指数的变化反映了不同类型群落在研究区内的镶嵌分布,从而形成了该区域内群落的多样性和复杂性.对植被样方与地形地貌因子的CCA排序结果显示,植被群落空间格局的形成更多地取决于区域内的水分变化梯度.在本文的研究尺度上,这种水分变化梯度主要由微地形地貌的变化来决定,而不是由与黄河的距离因子来决定.因此,在本区域的土地开发利用过程中,应充分考虑目前已经形成的相对稳定的地形地貌单元特点,在缺乏完善的风沙防护措施和排水除盐措施的情况下,不宜采取大规模平整土地进行综合开发.     

Distribution and reproductive success of the Pied Flycatcher Ficedula hypoleuca in relation to forest patch size and vegetation characteristics; the effect of scale

We studied habitat choice, distribution and reproductive success in the Pied Flycatcher Ficedula hypoleuca in relation to the spatial structure of a fragmented forest area. Two hierarchical levels of forest habitat structure were used: (1) forest patch size (macrohabitat) and (2) vegetative structure within forest patches (microhabitat). In spring, both males and females settled preferentially in large and medium-sized forest stands (>1 ha) where breeding density was also higher than in small stands (<1 ha). Stands <5 ha were occupied later, and the proportion of unpaired males was higher there than in large stands (>5 ha). The known age distributions of breeding birds and breeding success were independent of forest patch size. Nest predation rate was not associated with stand size or nest distance from the forest edge. The preference of breeding birds for large forest patches was presumably related to the higher amount of resources (nest sites, mates, food) a large patch can offer for reproduction compared with a small patch. At the microhabitat level, territories of old males were characterized by relatively more deciduous trees that contained more invertebrate food than coniferous tree-dominated territories of yearling males. The reproductive success of old males, as measured by the number of fledged young per male, was higher than that of yearling males. This suggests that the larger body size and blacker plumage of old males possibly contributed to male dominance and that old males excluded younger males from preferred habitats through territorial behaviour. Our results suggest that habitat choice of the Pied Flycatcher was affected by both micro- and macrohabitat. On the basis of settlement pattern, density and reproductive success, the distribution of Pied Flycatchers across forest stands of different sizes followed the ideal-free distribution model, whereas at the microhabitat level, age-related unequal distribution of males followed the ideal-despotic model of Fretwell and Lucas.     

不同干扰强度下针阔混交林树种空间格局及种间关联性

为了探究不同干扰程度下针阔混交林树种空间格局变化及树种种间关联性,本研究以关帝山庞泉沟自然保护区不同干扰强度的针阔混交林为对象,选取郁闭度、林分密度、伐桩数量作为划分干扰程度的因子,采用相对影响法,将林分划分为未干扰、中度干扰和严重干扰3个等级。利用空间点格局K2函数分析了林分的空间分布格局及其不同树种的种间关联。结果表明: 未干扰样地直径分布呈倒“J”型,中度和严重干扰样地直径分布均为双峰曲线分布。同时,未干扰和中度干扰林分在小尺度上呈现聚集分布,严重干扰的林分随机分布更为显著。针阔树种种间关系表现为未干扰林分在所有尺度上无关联;小尺度上中度干扰表现为正相关,严重干扰林分表现为负相关;较大尺度上中度与严重干扰林分均表现为不相关。说明林分干扰程度越严重,林下小径级林木数量越少,导致小尺度上种内聚集程度越低。同时,适当的干扰程度有利于林分内物种间对环境资源的协同利用。研究揭示了干扰对林分结构的影响,可为林分结构调整及抚育经营提供理论依据。     

吉林蛟河针阔混交林功能性状分布格局及其对地形因素的响应

森林群落的构建过程及其内在机制是生态学研究的热点问题。植物功能性状是指能够代表植物的生活史策略,反映植物对环境变化响应的一系列植物属性。通过植物功能性状的分布格局及其对环境因素的响应有助于推测群落的构建过程及其内在作用机制。以吉林蛟河21.12hm²温带针阔混交林样地为研究对象,采集并测量了样地内34种木本植物的6种不同的功能性状。以20m×20m的样方为研究单元,通过计算平均成对性状距离指数（mean pairwise trait distance;PW）和平均最近邻体性状距离指数（mean nearest neighbor trait distance;NN）来探讨群落中单个性状和综合性状的分布格局。同时结合地形因子采用回归分析探讨功能性状的分布格局对局域生境变化的响应。基于PW的结果显示：单个性状中除叶面积外,其余性状的分布格局均为聚集分布多于离散分布;基于NN的结果显示：除叶面积和最大树高外,其余性状的分布格局为聚集分布多于离散分布。此外,由6种单个性状组成的综合性状的分布格局同样为聚集分布多于离散分布。基于回归分析的结果显示：森林群落中功能性状的分布格局受到海拔、坡度和坡向等因素的显著影响,而凹凸度的影响则不显著。研究结果表明包括环境过滤和生物相互作用的非随机过程能够影响温带针阔混交林的群落构建过程,中性过程对该区域群落构建过程的影响不显著。     

Response of pine natural regeneration to small‐scale spatial variation in a managed Mediterranean mountain forest

Questions: What influence do management practices and previous tree and shrub stand structure have on the occurrence and development of natural regeneration of Pinus sylvestris in Mediterranean mountain forests? How are the fine‐scale and environmental patterns of resources affected and what impact does this have on the distribution of the regeneration? Location: A Pinus sylvestris Mediterranean mountain forest in central Spain. Methods: Upperstory trees and regeneration (seedlings and saplings) were mapped in four 0.5‐ha plots located in two types of stand with different management intensities (even‐aged and uneven‐aged stands). Environmental variables were recorded at the nodes of a grid within the plots. The relationships between the upperstory and regeneration were evaluated by bivariate point pattern analysis; redundancy analysis ordination and variation partitioning were performed to characterize regeneration niches and the importance of the spatial component. Results: Seedlings and saplings presented a clumped structure under both types of management and their distribution was found to be related to the spatial distribution of favourable microsites. Regeneration was positively related to conditions of partial cover with high soil water content during the summer. More than half of the explained variance was spatially structured in both types of stand. This percentage was particularly high in the even‐aged stands where the pattern of regeneration was highly influenced by the gaps created by harvesting. Conclusions: The spatial distribution of the tree and shrub upperstory strongly influences regeneration patterns of P. sylvestris. Current management practices, promoting small gaps, partial canopy cover and moderate shade in even‐aged stands, or favouring tree and shrub cover in the case of uneven‐aged stands, appears to provide suitable conditions for the natural regeneration of P. sylvestris in a Mediterranean climate.