A measure for spatial heterogeneity of a grassland vegetation based on the beta‐binomial distribution

Abstract. A method is proposed to estimate the frequency and the spatial heterogeneity of occurrence of individual plant species composing the community of a grassland or a plant community with a short height. The measure is based on the beta‐binomial distribution. The weighted average heterogeneity of all the species composing a community provides a measure of community‐level heterogeneity determining the spatial intricateness of community composition of existing species. As an example to illustrate the method, a sown grassland with grazing cows was analysed, on 102 quadrats of 50 cm × 50 cm, each of which divided into four small quadrats of 25 cm × 25 cm. The frequency of occurrence for all the species was recorded in each small quadrat. Good fits to the beta‐binomial series for most species of the community were obtained. These results indicate that (1) each species is distributed heterogeneously with respective spatial patterns, (2) the degree of heterogeneity is different from species to species, and (3) the beta‐binomial distribution can be applied for grassland communities. In most of the observed species spatial heterogeneity is often characterized by species‐specific propagating traits: seed‐propagating plant species exhibited a low heterogeneity/random pattern while clonal species exhibited a high heterogeneity/aggregated pattern. This measure can be applied to field surveys and to the estimation of community parameters for grassland diagnosis.     

禁牧条件下不同类型草地群落结构特征

利用幂乘方法则模型探讨了鄂尔多斯3种不同类型草地在禁牧情况下的群落结构特征,包括物种组成、物种多样性、生物量和空间分布规律。结果表明:幂乘方法则在解析鄂尔多斯不同类型草地的空间异质性时具有很好的吻合性;羊草(Leymuschinensis)草地、芨芨草(Achnatherum splendens)草地和油蒿(Artemisia ordosica)灌丛草地均比随机分布呈现了较强的空间异质性,群落整体的空间异质性指数表现为油蒿灌丛草地>羊草草地>芨芨草草地;群落整体的物种多样性指数为油蒿灌丛草地>羊草草地>芨芨草草地;L-样方(50cm×50cm)内的平均物种数和物种多样性指数均表现为羊草草地极显著地高于油蒿灌丛草地,油蒿灌丛草地又极显著地高于芨芨草草地(P<0.001);L-样方内的平均生物量表现为油蒿灌丛草地极显著地高于芨芨草草地(P<0.001),而羊草草地与油蒿灌丛草地以及芨芨草草地之间没有显著性差异;这3种类型的草地其L-样方内的平均生物量和物种多样性指数都随着群落整体空间异质性指数的增大而增大。     

Spatial pattern changes in aboveground plant biomass in a grazing pasture

Using gamma distribution and spatial autocorrelation, it was demonstrated that plant biomass per unit area of a pasture grazed by cattle exhibited two kinds of spatial heterogeneity: small-scale heterogeneity caused by grazing and large-scale heterogeneity caused by topography, land aspect, etc. For each of the 10 measurement times from May to August, 100 quadrats 50cm × 50cm were arranged along a straight line 50m long in a pasture, and the plants within the quadrats were harvested at the height of 3cm above the ground surface to measure the dry weight. The data were aggregated into frequency distributions, and gamma distribution and the parameter values were estimated. This analysis showed that with the progression of grazing the amount of biomass decreased and the degree of spatial heterogeneity in biomass, measured per 0.25m², increased, and due to plant regrowth the trends were reversed. By rearranging the 100 biomass data in order of weight, it was suggested that plots with an extremely large biomass were not grazed by cattle and remained in the pasture. For the same data, variations of biomass along the straight line were divided into two parts based on the moving average: the spatial trend and the residuals which cannot be explained by the trend. In this analysis, 48–75% of the total spatial variation was explained by the trend along the straight line. Analysis using spatial autocorrelation for the actual biomass changes showed that the biomass changes within a range of about 10m on the straight line gave a positive correlation, which indicates a topographical trend in biomass. Spatial autocorrelation for residuals suggested that the spatial changes in biomass along the straight line followed a wave-like or checker-board pattern. Small-scale spatial heterogeneity in plant biomass may be caused by the uneven deposition of excreta by grazing animals, uneven use of the grassland by grazing animals, and uneven dispersal of plant seeds through faeces over the grassland. The possibility that such unevenness might accelerate energy flow in the grassland ecosystem and contribute to grassland sustainability is discussed.     

Spatial pattern model of herbaceous plant mass at species level

Individual plant species distribute according to their own spatial pattern in a community. In this study, we proposed an index for measuring the spatial heterogeneity in mass (dry weight) of individual plant species. First, we showed that the frequency distributions for mass of individual plant species per quadrat in a plant community are expressed using the gamma distribution with two parameters of λ (mean) and p. The parameter p is a measure indicating the level of spatial heterogeneity of plant mass as follows: (1) when p = 1, the plant mass per quadrat has a random pattern; (2) when p > 1, the plant mass has a spatial pattern with a lower heterogeneity than would be expected in the random pattern; and (3) when p < 1, the plant mass has a spatial pattern with a higher heterogeneity than would be expected in the random pattern. The p value for a given species can easily be calculated by the following equation if we use the moment method: (mean plant mass among quadrats)² / (variance of plant mass among quadrats). The scatter diagram of (λ, p) for individual plant species, exhibits the spatial characteristics of each species in the community. We illustrated two examples of the (λ, p) diagram from data for individual species composing actual communities in a semi-natural grassland and a weedy grassland. Frequency distributions for the plant mass of individual species per quadrat followed the gamma distribution, and indi vidual species exhibited an inherent level of spatial heterogeneity.     

Analysis of Grassland Vegetation of the Southwest Heilongjiang Steppe (China) Using the Power Law

In 1997, we conducted a vegetation survey in three semi-arid natural grasslands (steppes) withdifferent livestock grazing intensities in Southwest Heilongjiang Province, China. The dominant grasslandspecies was the grass Stipa baicalensis Roshev. Grasslands with light, intermediate, and heavy grazingintensities were located 10, 5, and 2 km from a village, respectively. Villagers use the steppe to raise cattle,horses, sheep, and goats. Each of the three grasslands was surveyed by placing 100 quadrats (50 cm×50 cm)along a 50 m line transect. Each quadrat was divided into four equal areas (25 cm×25 cm; S-quadrats) and allplant species occurring in each of these smaller areas were identified and recorded. These data were summa-rized into frequency distributions and the percentage of S-quadrats containing a given species and thevariance of each species were estimated. The power law was applied to these estimates. The power law wasused to evaluate the spatial heterogeneity and frequency of occurrence for each species in the grasslandcommunity. The lightly grazed grassland exhibited high spatial heterogeneity (caused by large plant size),the highest species diversity, and a high occurrence of S. baicalensis. In contrast, the heavily grazedgrassland exhibited high spatial heterogeneity (caused by patchy populations of small plant size), lowspecies diversity, and a low occurrence of S. baicalensis. We judged that the heavily grazed grassland wasovergrazed and exclusion of livestock from the degraded areas is necessary for recovery.     

Grassland dynamics under sheep grazing in an Australian Mediterranean type climate

Grassland dynamics in a degraded disclimax grassland dominated by Danthonia caespitosa Gaudich. are examined using both demographic and multivariate approaches in an experiment designed to determine the effect of grazing intensity and exclosure on pasture dynamics. The experiment ran for 20 years from 1949 to 1968, using permanent quadrats at 3 grazing intensities and within exclosures. Demographic studies of some perennial grass species demonstrated markedly different responses to grazing; Danthonia caespitosa was unaffected by grazing but responsive to seasonal rainfall differences. Enteropogon acicularis survived only on protected sites. Numerical classification of total species set (121 species) for six observation periods demonstrated that community types were sensitive to differences in winter rainfall, and time since the start of experiment. Principal component analysis of permanent quadrat observations for individual years demonstrates quadrat trajectories which confirm this and indicate progressive divergence of the successional trends of the grazed and ungrazed quadrats. Repeated analysis on grazed quadrats only, shows that three components of pasture dynamics can be recognized; these are trend (succession?) and seasonal differences, each of which account for about 20% of the variance, and differences due to soil heterogeneity in the experimental paddock (8% of variance accounted for). No effect of grazing intensity was detected. Multivariate techniques can provide a clear partitioning of types of dynamic behaviour present in grassland communities. It is concluded that partitioning of environmental heterogeneity prior to demographic studies would increase their sensitivity.

     

Analysis of Grassland Vegetation of the Southwest Heilongjiang Steppe （China） Using the Power Law

In 1997, we conducted a vegetation survey in three semi-arid natural grasslands （steppes） with different livestock grazing intensities in Southwest Heilongjiang Province, China, The dominant grassland species was the grass Stipa baicalensis Roshev. Grasslands with light, intermediate, and heavy grazing intensities were located 10, 5, and 2 km from a village, respectively. Villagers use the steppe to raise cattle, horses, sheep, and goats. Each of the three grasslands was surveyed by placing 100 quadrats （50 cm×50 cm） along a 50 m line transect. Each quadrat was divided into four equal areas （25 cm×25 cm; S-quadrats） and all plant species occurring in each of these smaller areas were identified and recorded. These data were summarized into frequency distributions and the percentage of S-quadrats containing a given species and the variance of each species were estimated. The power law was applied to these estimates. The power law was used to evaluate the spatial heterogeneity and frequency of occurrence for each species in the grassland community. The lightly grazed grassland exhibited high spatial heterogeneity （caused by large plant size）, the highest species diversity, and a high occurrence of S. baicalensis. In contrast, the heavily grazed grassland exhibited high spatial heterogeneity （caused by patchy populations of small plant size）, low species diversity, and a low occurrence of S. baicalensis. We judged that the heavily grazed grassland was overgrazed and exclusion of livestock from the degraded areas is necessary for recovery.     

Checkerboard score–area relationships reveal spatial scales of plant community structure

Identifying the spatial scale at which particular mechanisms influence plant community assembly is crucial to understanding the mechanisms structuring communities. It has long been recognized that many elements of community structure are sensitive to area; however the majority of studies examining patterns of community structure use a single relatively small sampling area. As different assembly mechanisms likely cause patterns at different scales we investigate how plant species co‐occurrence patterns change with sampling unit scale. We use the checkerboard score as an index of species segregation, and examine species C‐score1–sampling area patterns in two ways. First, we show via numerical simulation that the C‐score–area relationship is necessarily hump shaped with respect to sample plot area. Second we examine empirical C‐score–area relationships in arctic tundra, grassland, boreal forest and tropical forest communities. The minimum sampling scale where species co‐occurrence patterns were significantly different from the null model expectation was at 0.1 m² in the tundra, 0.2 m² in grassland, and 0.2 ha in both the boreal and tropical forests. Species were most segregated in their co‐occurrence (maximum C‐score) at 0.3 m² in the tundra (0.54 3 0.54 m quadrats), 1.5 m² in the grassland (1.2 3 1.2 m quadrats), 0.26 ha in the tropical forest (71 3 71 m quadrats), and a maximum was not reached at the largest sampling scale of 1.4 ha in the boreal forest. The most important finding is that the dominant scales of community structure in these systems are large relative to plant body size, and hence we infer that the dominant mechanisms structuring these communities must be at similarly large scales. This provides a method for identifying the spatial scales at which communities are maximally structured; ecologists can use this information to develop hypotheses and experiments to test scale‐specific mechanisms that structure communities.     

A novel photographic approach for monitoring the structural heterogeneity and diversity of grassland ecosystems

Aims Studies that investigate the space-filling heterogeneity of biological structures in plant communities remain scarce. The main objective of this study was to evaluate the relationship between newly developed photographic measures of structural heterogeneity in digital images and plant species composition in the context of a long-term grassland experiment.Methods We tested a close-range photographic protocol using measures of structural heterogeneity in gray-tone images, namely mean information gain (MIG) and spatial anisotropy, to assess differences in the compositional (species richness) and functional characteristics (plant height and flowering) of 78 managed grassland communities. We also implemented a random placement model of community assembly to explore the links between our measures of structural complexity and the geometric pattern of plant communities.Important findings MIG and spatial anisotropy correlated with the growth and species richness of grassland communities. Simulations showed that structural heterogeneity in gray-tone digital images is a function of the size distribution and orientation pattern of plant modules. This easy, fast and non-destructive methodological approach could eventually serve to monitor the diversity and integrity of various ecosystems at different resolutions across space and time.     

Clustered animal burrows yield higher spatial heterogeneity

An understanding of the relationships between spatial heterogeneity and disturbance regime is important for establishing the mechanisms necessary to maintain biodiversity. Our objective was to examine how the configuration of disturbance by burrowing rodents (Siberian marmot) affected the spatial heterogeneity of vegetation and soil nutrient properties. We established three 2500-m² (50 m × 50 m) isolated-burrows plots and three 2500-m² clustered-burrows plots in a Mongolian grassland. Each plot was subdivided into 4-m² quadrats, and the plant species richness, percent coverage, and soil nutrient properties in the quadrats were surveyed. Spatial heterogeneity was calculated for vegetation using the mean dissimilarity of species composition among sample quadrats, and geostatistical analysis was used to calculate soil properties. Heterogeneous patches of plants such as Achnatherum splendens and higher nutrient concentrations were found only near the clustered burrows. As a result, spatial heterogeneities of vegetation and soil nutrient properties were higher in the clustered colony than those in the isolated colony. The configuration of disturbance patches affected the spatial heterogeneity at the landscape level through the spatial pattern of disturbance frequency.     

鄂尔多斯草地退化过程中个体分布格局与土壤元素异质性

应用复合个体-距离法分析了鄂尔多斯草地退化过程中多种群和优势种个体分布格局,应用统计方法分析了土壤有机碳和氮元素的异质化过程。结果表明：在鄂尔多斯草地退化过程中,多种群和优势种个体分布格局大多表现出不同尺度下的随机分布和聚集分布,但多种群的个体分布格局在本氏针茅(Stipa bungeana) 油蒿(Artemisia ordosica)群落中出现均匀分布;优势种油蒿和牛心朴子(Cymanchum komarovii)在油蒿 牛心朴子群落中均出现了均匀分布。土壤元素在单一优势种群落中存在不同程度的异质性,油蒿群落中异质化程度最强;在共优势种本氏针茅 油蒿群落中,出现了土壤元素相对均质化,油蒿 牛心朴子群落中,异质化程度相对减弱。由于对土壤水分和养分的竞争,使共优势种群落出现植物的均匀分布,土壤元素相对均质化。土壤元素先后异质化过程首先表现为土壤有机质的异质化过程,然后才表现为土壤氮的异质化过程。土壤元素先后异质化过程表明异质化过程首先表现为植被异质化,植被的异质化导致土壤某些元素的异质化。     

Positive diversity–invasibility relationship in species-rich semi-natural grassland at the neighbourhood scale

Background and Aims

Attempts to answer the old question of whether high diversity causes high invasion resistance have resulted in an invasion paradox: while large-scale studies often find a positive relationship between diversity and invasibility, small-scale experimental studies often find a negative relationship. Many of the small-scale studies are conducted in artificial communities of even-aged plants. Species in natural communities, however, do not represent one simultaneous cohort and occur at various levels of spatial aggregation at different scales. This study used natural patterns of diversity to assess the relationship between diversity and invasibility within a uniformly managed, semi-natural community.

Methods

In species-rich grassland, one seed of each of ten species was added to each of 50 contiguous 16 cm² quadrats within seven plots (8 × 100 cm). The emergence of these species was recorded in seven control plots, and establishment success was measured in relation to the species diversity of the resident vegetation at two spatial scales, quadrat (64 cm²) within plots (800 cm²) and between plots within the site (approx. 400 m²) over 46 months.

Key Results

Invader success was positively related to resident species diversity and richness over a range of 28–37 species per plot. This relationship emerged 7 months after seed addition and remained over time despite continuous mortality of invaders.

Conclusions

Biotic resistance to plant invasion may play only a sub-ordinate role in species-rich, semi-natural grassland. As possible alternative explanations for the positive diversity–invasibility relationship are not clear, it is recommended that future studies elaborate fine-scale environmental heterogeneity in resource supplies or potential resource flows from resident species to seedlings by means of soil biological networks established by arbuscular mycorrhizal fungi.     

Measures of spatial heterogeneity for species occurrence or disease incidence with finite-counts

We propose two types of indices with finite-count correction to measure the spatial heterogeneity of binary characteristics of organisms, such as occurrence or non-occurrence of organisms and infected or non-infected plants. We consider the following two examples: plant occurrence in a grassland community, and yellow dwarf disease infection in a rice field. For the grassland community, N quadrats comprising n cells of equal area, were set at random sites in a grassland, and the occurrence of a given species A in each of n cells was recorded. For disease infection, N quadrats, each consisting of n rice plants, were set at random sites in a paddy field, and the number of plants infected with yellow dwarf virus in each quadrat were counted. In these examples, since the number of cells in a quadrat is finite, neither occurrence nor incidence increase infinitely, unlike the number of aphids on a maize leaf. The first category of index belongs to the mean : variance ratio type. The estimated index value for occurrence (or incidence) is the same as that for non-occurrence (or non-incidence). The second category belongs to the k-type of a negative binomial distribution. If some random plants die or recover from the disease then the expected value of the second type of index does not change. For n, the current indices approach the mean : variance ratio and the inverse of k in a negative binomial distribution, respectively. This indicates that these indices are suitable for binary data sets.     

草原群落退化演替过程中微斑块土壤碳氮的空间异质动态

微斑块变化是草原退化过程中的活跃成分。分析了呼伦贝尔克氏针茅草原逆行演替过程中微斑块土壤全碳、全氮和碱解氮含量的空间异质性,提出了"养分聚集效应"的概念。研究结果表明:随着群落退化演替的加剧,土壤全碳、全氮和碱解氮的含量均表现为演替前期演替后期演替中期(P0.05)。从土壤全碳、全氮和碱解氮的变异系数和变异函数综合分析来看,10 cm×10 cm微尺度上,草原退化演替过程中土壤全碳、全氮和碱解氮的空间异质性具有明显的不一致性;全碳的空间异质性表现为演替中期演替前期演替后期,全氮表现为演替后期演替前期演替中期,碱解氮表现为演替中期演替后期演替前期。草原退化过程中土壤养分在微斑块上的富积和迁移表现出尺度依赖性和变异性。     

云南沾益中度退化天坑草地植物群落水平空间分布特征

规模最为宏大的地表负地形——喀斯特天坑不仅是世界罕见的地质奇观,圈闭化地形而形成的独特生境更是探索植物物种多样性及其空间分布特征最为理想的地域。以云南沾益天坑群中的中度退化天坑——"巴家陷塘"坑底草地植物群落为研究对象,通过α多样性指数与β相似性系数的应用,对"巴家陷塘"中度退化天坑坑底草地植物群落物种多样性的水平空间分布特征展开探讨,并对其近邻退化漏斗和"大毛寺"原生天坑坑外草地植物群落的相似性进行对比分析,研究结果发现:(1)中度退化天坑坑底草地植物群落物种丰富度和多样性从中心到边缘呈现出"中心高边缘低"的"逆洼地效应"特征;(2)相较于土壤资源,喀斯特天坑植物群落的生长与空间分布对光照资源的响应更为明显,且四周坑壁高度差异造成的坑内光照资源的空间异质性,也使坑底草地植物物种多样性存在一定的"圈层效应";(3)喀斯特天坑独特的圈闭化地形对植物群落的"隔离"、庇护作用即使在退化天坑中也有明显体现,且对喜阴、耐湿的物种更具庇护性。因而在全球气候变化下,即使是退化天坑,也应成为区域小尺度生态避难所的重要保护对象。     

Three-dimensional soil heterogeneity modulates responses of grassland mesocosms to an experimentally imposed drought extreme

Heterogeneity is an intrinsic characteristic of soils, which regulates plant diversity and ecosystem functioning. However, whether soil heterogeneity also modulates responses of plant communities to climate change, including climate extremes, remains largely an open question. Here, we explore responses of plant communities to drought extremes across four levels of spatial soil heterogeneity, with cell sizes varying from very small to very large, i.e. 0, 12, 24 and 48 cm. These were created in mesocosms by alternating nutrient-rich and nutrient-poor substrate in three dimensions. A seed mixture of 24 grassland species was evenly sown on each mesocosm in spring. In late summer, a three-week drought was imposed with a rainout shelter. During the drought, soil water content at the mesocosm scale decreased more at intermediate (12 and 24 cm) than at small or large (0 and 48 cm) cell sizes, which was reflected in increased senescence and drought-induced heat stress. These responses could be traced to greater plant biomass coupled with higher water demand at those intermediate cell sizes, likely related to between-cell access to nutrients and effects of diversity and community composition. Our results indicate that soil heterogeneity can modulate the impact of drought extremes on plant communities, though more research is needed on the transition between intermediate and extreme cell sizes, where heterogeneity effects seem to change most. We propose that soil heterogeneity be considered more explicitly in studies of changing precipitation regimes.     

Spatial heterogeneity of soil nutrients and plant species in herb-dominated communities of contrasting land use

Recent interest in spatial pattern in terrestrial ecosystems has come from an awareness of the intimate relationship between spatial heterogeneity of soil resources and maintenance of plant species diversity. Soil and vegetation can vary spatially in response to several state factors of the system. In this study, we examined fine-scale spatial variability of soil nutrients and vascular plant species in contrasting herb-dominated communities (a pasture and an old field) to determine degree of spatial dependence among soil variables and plant community characteristics within these communities by sampling at 1-m intervals. Each site was divided into 25 1-m² plots. Mineral soil was sampled (2-cm diameter, 5-cm depth) from each of four 0.25-m² quarters and combined into a single composite sample per plot. Soil organic matter was measured as loss-on-ignition. Extractable NH₄ and NO₃ were determined before and after laboratory incubation (28 days at 27°C) to determine potential net N mineralization and nitrification. Cations were analyzed using inductively coupled plasma emission spectrometry. Vegetation was assessed using estimated percent cover. Most soil and plant variables exhibited sharp contrasts between pasture and old-field sites, with the old field having significantly higher net N mineralization/nitrification, pH, Ca, Mg, Al, plant cover, and species diversity, richness, and evenness. Multiple regressions revealed that all plant variables (species diversity, richness, evenness, and cover) were significantly related to soil characteristics (available nitrogen, organic matter, moisture, pH, Ca, and Mg) in the pasture; in the old field only cover was significantly related to soil characteristics (organic matter and moisture). Both sites contrasted sharply with respect to spatial pattern of soil variables, with the old field exhibiting a higher degree of spatial dependence. These results demonstrate that land-use practices can exert profound influence on spatial heterogeneity of both soil properties and vegetation in herb-dominated communities.     

黄土丘陵沟壑区不同坡向撂荒草地植物群落种群空间格局

本研究以黄土丘陵沟壑区4种不同坡向的草地植物群落为研究对象,探讨其物种组成和多样性特征,并运用幂乘方法则分析了群落的空间异质性和种群空间格局.结果表明: 铁杆蒿在不同坡向均有分布,为群落优势种;其他物种在不同坡向群落中的地位及对群落空间格局的贡献不同.不同坡向植物群落的空间分布格局均为集群分布,且聚集程度大小为阳坡＞半阳坡＞半阴坡＞阴坡.群落空间异质性指数与Simpson多样性指数呈显著正相关,与Margalef丰富度指数、Shannon多样性指数和Pielou均匀度指数呈显著负相关.优势种铁杆蒿、亚优势种达乌里胡枝子和长芒草,以及少数伴生种如猪毛蒿、臭草的相对空间异质性指数大于群落整体空间异质性指数,使群落趋于集群分布;多数伴生种如芦苇、苦荬菜、白羊草的相对空间异质性指数与群落整体空间异质性指数相近,它们与群落自身分布状态一致;刺儿菜、米口袋、茭蒿等偶见种的相对空间异质性指数小于群落整体空间异质性指数,降低了群落的聚集程度.综上,群落的聚集程度主要由优势种、亚优势种和偶见种共同决定,其中优势种、亚优势种和部分伴生种促使群落呈聚集分布,偶见种则降低了群落的集群分布.     

内蒙古草原退化群落恢复演替的研究I. 退化草原的基本特征与恢复演替动力

 本项研究自1983年起在锡林河中游对放牧退化的冷蒿(Artemisia frigida)占优势的草原群落变型进行封育恢复实验与长期监测。每年在植物生长季以15天为间隔进行取样测定,即每年测定9期,每期做10或20个1m×1m的样方。测定项目包括：群落中各植物种群的地上现存生物量、密度、高度、花(果)枝数等。还采用改进的样方方差法监测植物种群空间分布格局的动态。并同时在保护良好的羊草+大针茅(Leymus chinensis+Stipa grandis)群落中进行测定取得完全对应的数据,作为对照系列。以上两种群落的土壤水分与养分动态的长期监测由本站土壤组承担。根据连续十二年监测数据的分析,对退化草原群落的性质与特征提出以下的认识,并对退化草原恢复演替的驱动因素进行厂探讨。1．草原退化演替阶段是与一定强度的放牧压力保持平衡而相对稳定的群落变形,退化阶段取决于牧压强度与持续的年代。2．当群落退化到冷蒿为主要优势种的阶段时,与原生群落的种类组成相比,只发生一定的数量消长变化,对群落的物种丰富度影响不大。3．退化群落植物种群空间格局的均匀性较高,随着恢复演替的进展,因一些种群斑块增大而使空间不均匀性增强。4．退化群落与其原生群落的种—生物量关系呈对数正态模式,其演替过渡阶段成为分割线段模式,也反映出群落资源分配格局与群落空间格局的关系。5．退化草原的显著特征是植被生产力下降,冷蒿群落的生物量下降到原生群落的30％～40％,家畜嗜食的植物种减少50％～70％总生产力不足原生群落的30％。6．退化群落在自然封育条件下能够迅速恢复的原因,可归结为植物在削除放牧干扰后的种群拓殖能力与群落资源(水分,矿质养分等)的剩余。群落资源条件是种群拓殖的物质基础,从而成为恢复演替的动力。     

High shoot plasticity favours plant coexistence in herbaceous vegetation

Several theoretical considerations imply that high shoot morphological plasticity could increase competition symmetry and favour plant coexistence. We tested whether mean plasticity across co-occurring species is a key trait for explaining ramet density and species richness in herbaceous vegetation. We used three data sets to test the hypotheses: (a) experimentally achieved estimates of plasticity to light availability for 35 herbaceous species; (b) richness, ramet density and canopy architecture data from 17 herbaceous communities; (c) species richness data from a 5-year permanent-plot study in a calcareous grassland. In herbaceous communities containing species with relatively higher shoot plasticity, ramet density was significantly higher. Consequently, relatively more species were growing per unit area—a greater proportion of the community species pool was represented on 1 m². In the permanent plot study species-richness was higher in those 40×40 cm quadrats where species with high shoot plasticity prevailed—there was a positive regression of richness on the mean plasticity of species. This relationship was highly significant in five consecutive years. Our results suggest that shoot plasticity to light availability is evidently one of the key traits in processes that alter the density of co-existing plants and, therefore, species diversity in herbaceous communities. Electronic Supplementary Material Supplementary material is available for this article at