Fine‐scale five‐year vegetation change in boreal bog vegetation

Abstract. Within an ombrogenous part of N. Kisselbergmosen, Rødenes, SE Norway, fine‐scale changes in species abundance, successional trends relative to the main gradients (as represented by DCA axes), and co‐ordinated change within pairs of the bottom layer species are studied. Data sets were sampled twice with a five‐year interval, and included species abundance and cover of mud bottom, naked peat and litter in 436 sample plots (16 cm× 16 cm), and species abundance in 6976 subplots (4 cm× 4 cm). Depth from the surface of subplots to the water table was estimated in 1991. Most summers and growing seasons were somewhat drier than normal in the 5‐yr period. The area covered by mud‐bottom, naked peat and litter increased significantly, as did the frequencies of the dwarf shrubs Calluna vulgaris and Andromeda polifolia in hummocks and upper lawn. Sample plots were significantly displaced downward the peat productivity gradient (DCA 2), reflecting the reduced cover of many bottom layer species, including all Sphagnum spp. Significant coordinated changes in cover of bottom layer species are described. The changes observed in hummocks support the existence of a local regeneration cycle, as suggested by other researchers. Some of the vegetation changes seem parallel to those reported from areas with a higher nitrogen deposition, but it is not likely that nitrogen deposition alone is the major cause of the observed changes. Between‐year variation in population size and climatic fluctuations may as well explain the observed changes.     

Succession of terrestrial macrofungi along a deglaciation gradient at Glacier Blåisen, South Norway

A survey of terrestrial macrofungi along a deglaciation gradient from the glacier forefront to the established vegetation has been performed. The study was accomplished along a transect from Lake Finsevann to Glacier Blåisen, South Norway. Fortysix 6 m × 6 m plots divided into 9 subplots were positioned along this transect. The frequency of fruit bodies of each species in each plot was given on a 0–9 scale reflecting their occurrence in the subplots. Biotic and abiotic environmental variables for the plots were also recorded. The numerical methods used include principal component analysis (PCA), detrended correspondence analysis (DCA), and canonical correspondence analysis (CCA). Totally 99 species of macrofungi were recorded; 80.8 % of the species belonged to the Basidiomycota and 19.2 % to the Ascomycota. About one third of the species was only found once. In general, the species richness was highest on the glacier forefront, lowest in the most established vegetation. The proportion of species of Ascomycota was highest close to the glacier front. The main gradient in the data set was shown related to the primary ecological succession following the retreat of glaciers. The second gradient was possibly related to fine scale variation due to local moisture, microtopography, and secondary distribution. Of the variation in composition of species, 3 % can be ascribed to abiotic variance, 10 % to vascular plant variance, and 16 % to the combination of the abiotic and vascular plant component of variation. The results are discussed in the light of the time since deglaciation and the influence of certain biotic and abiotic environmental factors.     

Reseating of ecological gradients. III. The effect of scale and niche breath measurements

Abundances of eleven Sphagnum species in 800 sample plots are used to investigate the effect of DCA rescaling on Levins' measure of niche breadth relative to three partitions of the water-table gradient in a boreal Norwegian mire: (1) sample plots classified into 15 categories, each spanning an interval of 2 cm vertical extent, (2) as (1), but sequence of categories rescaled by DCA and sample plots reorganized into 15 categories with uniform beta diversity, and (3), sample plots ordinated by DCA and classified into 15 categories with uniform beta diversity by subdivision of an ordination axis highly correlated with median water-table. Habitat niche breadth is shown to be dependent on four issues (in order of supposedly decreasing importance): (1) scale, (2) noise level of data, (3) homogeneity of individual samples, and (4), weighting function. Six problems relevant to interpretation of measurements of niche breadth are discussed: (1) range of measures, (2) spacing of categories, (3) scale, (4) choice of gradients, (5) number of samples, and (6), comparability of studies. For measures of habitat niche breadth to be biologically meaningful, four conditions have to be satisfied: (1) the gradients studied have important impact on the studied species, (2) sampling is adequate, (3) scaling of gradients is in compositional turnover, and (4), comparability is demonstrated prior to comparison with other studies. Revisions of current methods are proposed. The role of DCA in niche studies is particularly emphasized.     

On the relationship between sample plot size and beta diversity in boreal coniferous forests

Nested sample plots of three sizes (16, 1, and 1/16 sq. m) from three different studies of Norwegian coniferous forests have been subjected to DCA ordination using the same choice of options. At each sample plot size, species quantities are recorded as frequency in 16 subplots. Beta diversity, measured as length of the first DCA axis, invariably increased upon lowering of sample plot size. The same applied to the eigenvalues of the axes. This is explained as a consequence of the weakening of structure in the data matrices when the fine-grained patterns of the vegetation are emphasized.     

An ecological approach to the investigation of a beech forest in Vestfold, SE Norway

The vegetation in a beech forest, Fritzøehusparken, is analysed using one hundred sample plots, each 25 m², distributed by a restricted random procedure. Percentage cover and frequency in subplots are used for quantification of species amounts, and their relative performance are evaluated. Advantages of choosing frequency in subplots are discussed. Thirteen environmental variables are measured from each of the sample plots. The vegetation is divided into four topographical/ecological types. This classification served as a reference in the further treatment. Ecological gradients corresponding to the most important vegetational gradients in the beech forest are identified by means of DCA ordination followed by statistical interpretation. Two main complex gradients were recognized; (1) the gradient in nutrient conditions, and (2), the gradient inlitter-wind conditions. The advantages of using DCA and statistical interpretation with integration of ecological measurements are emphasized.     

Persistence of vascular plants in a Norwegian boreal coniferous forest

Persistence, the tendency of a species to remain in its original position and not to colonize new sites, is studied for the most abundant forest vascular plants (25 species in spruce forest and 7 in pine forest) in Solhomfjell, Gjerstad, S Norway Data sets included presence/absence in 199 meso plots (1 m²) and 3184 subplots (1/16 m²), analyzed over a 5 yr interval, and a subset of 50 meso plots and 800 subplots, analyzed for six consecutive years Relationships between species variables (seedling frequency and mobility rate compiled from the literature, and cover and abundance means in the study area) are studied, and related to species optima along ecologically interpreted DCA ordination axes Vascular plant mobility may increase towards nutrient-poor sites Dominance in the boreal forest floor is mostly by clonal species Persistence was calculated for different temporal (1-5 yr) and spatial (1/16 and I m²) scales Persistence patterns in the spruce and pine forests were similar, but persistence decreased towards the xeric pine forests One main component of variation in persistence was demonstrated by PCA analyses the absolute level of persistence, which is related to seedling recruitment vs clonal growth, and within clonal species to ramet longevity, abundance, mobility, growth pattern and mode of surviving the infavourable season Minor components of variation in persistence were related to spatial scale and temporal scale Persistence characteristics were species-specific and varied little between years Numerous species characteristics were relevant to interpretation of variation in persistence, indicating a continuous, multidimensional variation in life history characteristics     

Pteridophyte diversity and species composition in four Amazonian rain forests

Abstract. Local variation in individual density, species composition, species richness and species diversity of terrestrial pteridophytes were studied at four sites in the tropical lowland rain forest of western Amazonia. 15 568 pteridophyte individuals representing 40 species were recorded in four plots. The variability among subplots within the same plot was considerable in all the characteristics measured (number of individuals, number of species, species diversity); the square 1‐ha plot was more homogeneous in these respects than any of the three 5 m by 1300 m transects. Species richness was affected by the density of individuals both within and among plots. Density of individuals was not affected by topographical position within any of the plots, whereas in some of the plots both species richness and species diversity were. Clustering and ordination analyses showed that floristically similar subplots could be found in different plots: although there was a tendency for subplots from the same plot to be floristically similar and therefore to group together, many recognized groups included subplots from two or more plots. Both within and among plots, the floristic differences corresponded to topographic position and were probably related to soil drainage. This was also evident in that the abundance patterns of many species followed the topography.     

海南岛霸王岭热带季雨林树木的死亡率

树木死亡是调节群落组成和结构的一种方式, 在森林生态系统动态中发挥着重要作用。该文在对海南岛霸王岭热带季雨林群落大量调查和树种功能群划分研究的基础上, 探索了热带季雨林群落及其不同功能群树木的死亡率及其随径级和环境条件的变化规律。结果表明: 海南岛霸王岭热带季雨林群落内树木死亡率的变化范围为3.42%-18.71%, 平均值为7.60%。按照功能群比较, 乔木死亡率显著高于灌木, 落叶树种死亡率高于常绿树种, 但具刺树种死亡率显著低于无刺树种。按照树木径级划分, 5-30 cm径级范围的死亡率均超过10%, 最高值出现在5-10 cm径级范围, 在相同径级范围内比较时, 落叶树种在 胸径(DBH ) < 5 cm时的死亡率显著高于常绿树种, 而在其他径级范围内, 二者之间无显著差异, 具刺和无刺树种的死亡率在所有径级范围内均无显著差异。乔木死亡率与群落距河流距离具有显著的相关性, 距离河流越远死亡率越高; 落叶树种和无刺树种则与坡位存在显著的相关性, 坡上部的死亡率显著高于坡中部。热带季雨林内树木的死亡率与其所处的群落生境有关, 较干旱的生境是导致树木死亡率较高的重要原因。     

Reseating of ecological gradients. II. The effect of scale on symmetry of species response curves

Response curves are presented for 15 species of vascular plants, bryophytes, and lichens relative to the water-table gradient in a boreal Norwegian mire. The gradient is scaled in two ways; position of median water-table in cm below the surface of the bottom layer, and in units of compositional turnover (rescaled by DCA). The two scalings produce quite different curves above the maximum water-level, which corresponds to median water-level 10–14 cm below the surface of the bottom layer. The significant drop in compositional turnover above this level can be ascribed to vertical differences in water-table having a lesser effect on plants in aerated hummocks than on plants in temporarily water-logged hollows. Skewness of response curves was reduced by gradient rescaling, most strongly after removal of influence of other gradients. It is argued that response curves will generally be Gaussian if: (1) the response is with respect to a dominant ecological factor, (2) the range of variation along the gradient is sufficient, (3) the distribution of samples is adequate, and (4), the gradient is scaled in units of compositional turnover.     

Plant niches along the water-table gradient on an ombrotrophic mire expanse

The vegetation within an ombrotrophic mire expanse m SE Norway is studied m detail Presence/absence of species and depth to the water table are recorded for 6976 plots (4x4 cm) The cumulative probability distributions along the water-table gradient are calculated for 24 species, and Inter-transect variability tested Shifts in realised depth to the water table (niche shifts) are frequent for most species It is concluded that interactions are of minor importance for species in the field layer The Sphagnum spp possess some attributes indicating equilibrium, while others indicate ongoing competition The patchy distribution of hepatics and lichens depend on chance and successional stage, and coexistence within these guilds depend on creation of new naked peat patches     

Plant composition in the Maya Biosphere Reserve: natural and anthropogenic influences

We studied the floristic composition and stand structure of lowland forests of the Maya Biosphere Reserve (MBR) of El Petén, Guatemala. Sampling was performed over a gradient of human influence, including a newly established returnee community (migrant), the cooperative Unión Maya Itzá, as well as an adjacent protected area within the core area of the MBR. Five 1-ha plots, each divided into 100 contiguous 10 × 10 m² subplots, were used to record presence of all tree and vine species. The study area is a low-diversity rainforest with a canopy layer dominated by species of Fabaceae, a shrub layer dominated by Rubiaceae, and vines dominated by Bignoniaceae. Vines were conspicuous both in number and in diversity. Variation in both tree life-form composition and canopy abundance pattern apparently reflect variation in intensity of forestry among the plots. The similarities between the plots in the cooperative (a logging plot and a settlement plot) with regard to species area curves, and mean number of tree and understory species per subplot, may indicate effects of disturbance (in general) on vegetation structure. Detrended correspondence analysis (DCA) was used to investigate gradients in species composition among the five plots (125 subplots) and to generate hypotheses about vegetation–environment relationships. The study area appears as a mosaic of site-specific forest types or associations determined by a particular species or groups of co-dominant species. The main environmental characteristics of the area determining species composition and structure are related to the drainage of soils and human disturbance.     

Seeding tallgrass prairie in monospecific patches promotes native species establishment and cover

In tallgrass prairie reconstruction, the way desired seeds are arranged on the landscape may affect species establishment, species persistence, and the establishment and persistence of undesired (nonseeded) species from the local propagule pool. To test effects of species seeding pattern on how grasslands develop spatially, we seeded 20—4 × 4–m bare soil plots with 16 tallgrass prairie species. Treatment plots were divided into 16—1 × 1–m subplots, 64—0.5 × 0.5–m subplots, 256—0.25 × 0.25–m subplots, or 1,024—0.125 × 0.125–m subplots. Each species was hand broadcast into separate subplots (1 m² total area/species) within each plot. An additional treatment included uniformly mixing and broadcasting all seeds across a plot. We recorded species cover at the 0.125 × 0.125–m scale within each plot at the beginning of the second and third growing seasons. While species persistence was greatest within plots seeded with larger subplots, plots with smaller subplots were more spatially diverse and less occupied by nonseeded species over time than larger subplot and mixed plots. As is common in reconstruction efforts, establishment was variable among species and seeding with monospecific subplots enhanced colonization of desired rhizomatous species (e.g., Heliopsis helianthoides, Monarda fistulosa, Elymus virginicus) into unoccupied locations at the expense of species from the local propagule pool. Results suggest that seeding species in smaller, monospecific patches could result in grasslands with a more balanced native species composition than those established with a seed mixture approach.     

哀牢山中山湿性常绿阔叶林木质藤本对边缘效应的响应

全球范围内森林片断化现象日益严重。与其他木本植物(乔木和灌木)相比, 木质藤本更趋向于分布在片断化森林的边缘, 因而了解木质藤本对边缘效应的响应对于进一步了解其对森林动态的影响极其必要。本文对哀牢山中山湿性常绿阔叶林林缘到林内环境梯度上木质藤本的变化进行了调查。在形成年龄分别为13年、35年和53年的3种类型的林缘, 设置从林缘向林内连续延伸的长方形样地(20 m × 50 m)各10个(总面积3 ha), 每个样地再划分为5个20 m × 10 m的样方。在每个样方内对胸径≥ 0.2 cm且长度≥ 2.0 m的木质藤本进行了每木调查。在3 ha的林缘样地中共记录到木质藤本植物2,426株, 隶属于14科19属31种。木质藤本的物种丰富度和多度均随距林缘距离的增加而降低, 边缘效应深度在35年林缘的边缘为30 m, 13和53年林缘的边缘则为20 m; 它们的胸高断面积在53年林缘的边缘效应深度为20 m, 但在13和35年林缘的不同距离上差异不显著。木质藤本对边缘效应的响应在物种水平上存在显著差异, 主要呈现正向和中性的响应格局, 包括只分布于林缘的物种, 和从林缘到林内环境梯度上密度逐渐降低的物种; 也有对边缘效应不敏感的物种。典范对应分析(CCA)表明, 林冠开度、边缘形成年龄和土壤水分是决定木质藤本在片断化森林边缘分布的重要影响因子。     

Shape matters in sampling plant diversity: Evidence from the field

The identification of shape and size of sampling units that maximises the number of plant species recorded in multiscale sampling designs has major implications in conservation planning and monitoring actions. In this paper we tested the effect of three sampling shapes (rectangles, squared, and randomly shaped sampling units) on the number of recorded species. We used a large dataset derived from the network of protected areas in the Siena Province, Italy. This dataset is composed of plant species occurrence data recorded from 604 plots (10 m × 10 m), each divided in a grid of 16 contiguous subplot units (2.5 m × 2.5 m). Moreover, we evaluated the effect of plot orientation along the main environmental gradient, to examine how the selection of plot orientation (when elongated plots are used) influences the number of species collected. In total, 1041 plant species were recorded from the study plots. A significantly higher species richness was recorded by the random arrangement of 4 subplots within each plot in comparison to the ‘rectangle’ and ‘square’ shapes. Although the rectangular shape captured a significant larger number of species than squared ones, plot orientation along the main environmental gradient did not show a systematic effect on the number of recorded species. We concluded that the choice of whether or not using elongated (rectangular) versus squared plots should dependent upon the objectives of the specific survey with squared plots being more suitable for assessing species composition of more homogeneous vegetation units and rectangular plots being more suited for recording more species in the pooled sample of a large area.     

Soil heterogeneity effects on canopy structure and composition during early succession

Early succession from annuals to herbaceous and woody perennials on abandoned fields and grasslands is marked by vegetation heterogeneity; i.e., variation in canopy composition and structure over the field through time. Soil resource heterogeneity could promote vegetation heterogeneity. We created soil resource patchiness on two newly ploughed fields by establishing two types (fertilized, unfertilized) of checkerboard plots with two, four, or eight subplots (trenched, untrenched) to test effects of soil fertility heterogeneity scale on vegetation heterogeneity during early succession. Canopy composition and structure were censused in years 2, 4 and 6. On both fields, soil resource heterogeneity did not affect canopy composition or structure. Differential abundance of dominant species, Ambrosia trifida on one field and Solidago altissima on the other, between fertilized and unfertilized plots led to decreased vegetation heterogeneity among subplots within plots and increased vegetation heterogeneity among plots. Soil enrichment promoted taller and layered, but species-poor, canopy. In general, soil fertility effects on abundance, expansion, and dominance of individuals of dominant species promoted variation in canopy composition and structure, or vegetation heterogeneity, among plots on the two fields.     

古田山中亚热带常绿阔叶林群落物种多样性的空间变异特征

古田山常绿阔叶林的群落组成、结构及其维持机制已有许多研究, 但该地区亚热带常绿阔叶林生物多样性空间变异特征还缺乏认识。本文以古田山24 ha大样地(划分为24个1 ha小样地)为基础, 具体分析了α多样性和β多样性在1 ha尺度上的空间变异特征。结果表明: (1)群落第一、二优势物种在各小样地之间变化不大, 但第三优势种变化较大; (2) α多样性变化中, 样地间木本植物个体数量变异最大, 物种丰富度其次, Pielou均匀度指数变异性最小; (3)物种丰富度与植株个体数量、Pielou均匀度指数没有显著的相关性, 与Shannon-Wiener指数呈显著正相关; Shannon-Wiener指数与Pielou均匀度指数呈显著正相关; (4)相邻样地间物种替代速率空间变异较大, 与物种丰富度的空间变化格局有明显差异。这些结果说明尺度对认识群落结构、探讨群落维持机制有重要作用; 由于森林群落是多尺度生态过程作用的结果, 大尺度样地可能有利于更好地揭示森林群落维持机制。     

Fine‐scale persistence of boreal bog plants

Abstract. Persistence, or the tendency of a species to remain in its original position without colonizing new sites, is studied for 24 species on the ombrotrophic Northern Kisselbergmossen in SE Norway. Data sets comprise presence/absence in 436 sample plots (16 cm x 16 cm) and 6976 subplots (4 cm x 4 cm) analysed with a 5‐yr interval. Persistence was calculated for both spatial scales, and the observed values were compared with null models of completely random presence/absence of species. Species characteristics (plot occurrences and persistence) were related to depth to the water table and species optima along ecologically interpreted DCA ordination axes. The observed persistence was significantly higher than predicted from the random model for all vascular plants and cryptogams at both spatial scales. All taxonomic groups showed higher persistence at the sample plot scale than at the subplot scale. No general relationship between persistence and depth to the water table was found, but for the cryptogams there was somewhat higher persistence for the less peat‐producing species. The persistence of the vascular plants depended on ramet longevity, growth form and vegetative mobility. In general, the observed persistence of most cryptogams was high, probably because of their perenniality, low growth rates and high reproductive output. Differences in growth‐form and life history, as well as the higher number of occurrences, are the most likely reasons for somewhat higher mean persistence of hepatics and Sphagna than of vascular plants at the subplot scale.     

海南岛霸王岭热带低地雨林植被恢复动态

热带次生林具有重要的物种保育和固碳功能, 然而高强度的干扰会导致次生林早期出现类似季雨林的阶段, 因而群落恢复速度和方向是当前热带生态学研究中最为关注的议题之一。该文以海南岛在刀耕火种弃耕地形成的不同演替阶段的次生林为研究对象, 比较森林不同恢复时间(12年、25年、55年)群落中的不同年龄(幼树、小树、成年树)个体与低地雨林老龄林的物种组成、多样性和群落结构差异, 探讨刀耕火种弃耕地恢复过程中的群落组配过程。首先, 在海南岛霸王岭林区内建立7个1 hm²(100 m × 100 m)的样地, 并调查样地内所有胸径≥ 1 cm的木本植物个体(不包括木质藤本)的种类、胸径大小和树高。无度量多维标定法(NMS)排序结果表明, 刀耕火种弃耕地恢复群落与老龄林的物种组成存在明显差异, 并且其物种组成差异随着径级增加而逐渐减小。刀耕火种弃耕地群落物种累积速度缓慢, 25年和55年恢复群落的种面积、种个体和种多度曲线无差异, 存在一个明显的停滞阶段。与物种组成相比, 群落结构恢复相对较迅速, 但仍没有形成老龄林阶段中的复杂结构。萌生个体在早期恢复群落中占有较高比例, 其个体密度和胸高断面积分别占总数的39.9%和55.9%, 但在恢复中后期迅速降低。刀耕火种弃耕地恢复群落中以先锋种和非先锋喜光种为主。虽然耐阴种随演替而逐渐增加, 但恢复中后期群落中的耐阴种重要值仅为老龄林的27.7%。这些结果表明, 虽然刀耕火种弃耕地恢复群落缓慢地逐渐接近最终恢复目标, 但仍然存在 很大的不确定性。刀耕火种弃耕地恢复过程中的异速恢复和停滞阶段需要纳入今后群落演替模型构建和森林固碳效益核甘共苦算中。     

Primary lowland dipterocarp forest at Danum Valley, Sabah, Malaysia. Species composition and patterns in the understorey

The spatial distributions of species of tree 10 cm gbh were examined in two 4 ha plots and related to the local variation in topography and soil chemistry. The plots were similar in their species composition, particularly in terms of the densities of small trees, and they showed very similar edaphic characteristics. Size class distributions varied little within and between plots. Ordination of 0.25 ha subplots highlighted parallel gradients in the vegetation of both plots when the densities of trees 10 cm gbh were considered. Focusing on understorey trees in the 10-<50 cm gbh class at the 0.04 ha subplot scale showed a similar vegetation gradient in both plots closely associated with change from lower slope to ridge. No relationship with soil chemistry was found. On the ridges a special group of understorey species formed clumps and these species contributed importantly to the ordinations. Borneo has a regional history of occasionally severe droughts. It is suggested here that the observed patterns in the understorey are due to differential responses to low soil water supply, the ridges probably tending to dryness more than the lower slopes. Within the large and diverse family Euphorbiaceae, which dominates the understorey at Danum, there may be ecophysiological groupings of species. The long-term effects of disturbance interacting with local edaphic factors on forest structure and composition are discussed.     

Small-scale variability as a mechanism for large-scale stability in mountain grasslands

Abstract. Data from a 7-yr permanent plot study of grassland dynamics were used to address the relationship between processes at two levels of resolution (3.3 cm x 3.3 cm, 50 cm x 50 cm). Grasslands mown and manured in the traditional way in the Krkonose Mts. (Riesengebirge) were used as a model system. Spatial dynamics at the finest scale were very high, as demonstrated by turnover of individual species in 3.3 cm x 3.3 cm subplots and year-to-year transition matrices of the same subplots. The direction of these dynamics was not correlated with grassland treatment, although there was some correlation within years. An extrapolation of such year-to-year dynamics to larger time scales would result in big large-scale changes on the community level, and large shifts in species composition of the whole sward. However, dynamics at larger spatial or temporal scales were generally small. Some directional change occurred in manured plots, whereas little change occurred in unmanured plots. Large-scale dynamics were not correlated with small-scale dynamics in plots without manuring, but some correlation was detectable in manured plots. There are probably several processes that drive small-scale dynamics, such as non-linear interactions and environmental fluctuations. We argue that within certain limits these forces act on species composition so as to make small-scale dynamics non-directional. This results in both large-scale species diversity and apparent large-scale stability of these grasslands. However, if these forces are beyond these limits, the small-scale dynamics may become directional, resulting in rapid changes at larger spatial scales.