Vegetation succession on the dunes near Oostvoorne,The Netherlands,since 1934, interpreted from air photographs and vegetation maps

The vegetation succession on the dunes near Oostvoorne, The Netherlands has been followed by means of a novel combination of repeated large-scale vegetation mapping and air photograph interpretation. Vegetation units have been discerned on the formation level because these could be distinguished fairly easily on the photographs and because the rates of change are appropriate to the time interval chosen. Nineteen formations were distinguished. Five 1:6250 maps were constructed, reflecting the formation pattern in 1934, 1943, 1959, 1972 and 1980. An overlay with 2736 grid points at 25 m field distance was used to quantify changes in the formation pattern.The results suggest a pronounced multiple pathway succession with nevertheless three principal trajectories of succession from pioneer to woodland vegetation. The outer dunes, which have originated since 1910, are distinct in successional characteristics from the inner dunes, which already existed but were released from heavy grazing pressure in 1910. The rate of change in the outer dunes was high in the beginning and is slowly decreasing eversince. In the inner dunes it went the other way around. Through visual extrapolation the likely formation patterns in 1910 and in 2000 were estimated.Transition frequencies proved highly variable for most formations. Moreover, strong spatial dependence was found. Limitations in the use of Markov models in cases of long-term succession in heterogeneous environments are discussed.Nomenclature follows the same sources as in van der Maarel et al. (1984).Field work carried out 1980–1981 when the authors were at the Division of Geobotany, University of Nijmegen. We thank Jos Rijntjes, Nijmegen, for his cooperation in the field. Vegetation maps were prepared and calculations performed at Uppsala. We thank the Foundation Het Zuid-Hollands Landschap, Rotterdam, for providing facilities and a grant for fieldwork as well as additional means to reproduce the vegetation maps. Two reviewers gave useful comments.     

Vegetation differentiation and secondary succession on a limestone hill in southern Poland

Abstract. The vegetation of the SkoHzanka Reserve, situated on a limestone hill near Krakow, was examined over a 30 year period, using the Braun-Blanquet method. With the help of a numerical classification eight types of grassland communities were distinguished, belonging to the classes Sedo-Scleranthetea, Nardo-Callunetea and Festuco-Brometea, and 11 types of woodland communities, both remnants of ancient woodlands from the classes Querco-Fagetea and Vaccinio-Piceetea and more recent woods. In the case of grasslands the results of detrended correspondence analysis (DCA) indicated that the largest variation was associated with the soil type. Woods on sandy soils are very different from those on calcareous soils, and the younger recent woods are very different from the ancient woodlands. An analysis of historical cartographic material and aerial photographs shows that in the last few decades the area of recent woods has increased considerably at the expense of grasslands. Changes in species diversity in the woods during secondary succession and their successional convergence were examined and discussed. It was found that young and mature woods on sandy soils are more similar to each other than young and mature woods on calcareous soils. These differences are mainly related to the fact that many woodland herb species growing in rich, broadleaved ancient woods are unable to colonize recent woods since they have a limited colonization capacity.     

云南大山包黑颈鹤自然保护区草甸植被的数量分类和排序

在法瑞学派样地调查法的基础上,采用双向指示种分析（TWINSPAN）和趋势对应分析（DCA）对大山包黑颈鹤自然保护区草甸植被进行数量分类和排序研究。调查样地包含114个物种,分属31科72属,其中禾本科（14种）、菊科（13种）、蓼科（8种）、石竹科（6种）、玄参科（6种）等科所含物种较多。经TWINSPAN分类,106个样地分为20个聚组,根据其指示种确定为20个群丛。样地DCA和物种DCA排序结果反映了植物群丛与指示种之间的一致关系,排序轴的生态学意义说明水分和海拔两个环境因子同时影响该区草甸植被的分布。     

广西阳朔岩溶植被演替过程种群变化及物种多样性

采用时空替代法研究广西阳朔岩溶植被在演替过程中的种群变化和物种多样性。研究表明,随着演替的进展,耐荫种群增加,乔木种群增加,优势种的更替呈现出一定的连续性,后一演替阶段的优势种往往已经隐含在前一演替阶段中;物种丰富度随演替进展表现出草丛<落叶阔叶林<灌丛<常绿落叶阔叶混交林的趋势;灌木层的物种丰富度在各演替阶段中均是最高的;在各演替阶段中,落叶阔叶林阶段的群落均匀度均表现为最大。     

Changes in soil and vegetation following stabilisation of dunes in the southeastern fringe of the Tengger Desert, China

Properties of the soil and sand-binding vegetation were measured at five sites plus a control on dunes of the Tengger Desert stabilized for periods of up to 50 years. In the topsoil, fine particles, total N, P, K and organic matter increased significantly with increasing site age. However, there were no significant changes in deeper soil profiles (>0.4 m depth). Soil pH, calcium carbonate content, and total salt content tended to increase with age. Soil water in the topsoil changed little with increasing age, but was closely related to rainfall during the 50-year period. For deeper soil layers (0.4–3.0 m) soil water decreased significantly with age. After revegetation, the number of herbaceous species increased up to 30 years and then levelled off to 12–14 species, whereas the number of shrub species decreased from the 10 initial sand-binding species to only 3 species. Shrub cover decreased from a highest average of about 33% to the current 9%, whereas cover and biomass of herbaceous species increased throughout succession from 1956 to 2006. The development of soil and cryptogamic crusts on the surface of stabilized dunes enhanced the colonization and establishment of herbaceous plants due to increasing water availability, clay and silt content and soil nutrients. We propose that changes in properties of the surface soil led to increased interception of water, favoring shallow rooted grasses and forbs over perennial shrubs.     

Vegetation development after the exclusion of grazing cattle in a meadow area in the south of Sweden

A study of 24 years of secondary succession was based on data from semi-permanent quadrats from investigations in 1952, 1955, 1968 and 1976, involving two exclosures, the first in a meadow grazed for hundreds of years but now abandoned, the second in a meadow mowed for hundreds of years and grazed for the last 50 years. A first order classification of quadrats produced units, which formed distinct spatial patterns indicating similar gradients, but also differences in response to the ceased grazing, in the two exclosures. A second order classification of units into groups revealed a rather simple structure of spatial and temporal relations. Eleven groups of species with similar behaviour could be recognized within a system of spatial and temporal species distributions. The vegetation in both exclosures developed towards an increased differentiation and heterogeneity or patchiness. The border between two soil types was clearly reflected in the spatial pattern of units. Rates of change were greatest in the beginning and were shown to closely follow logarithmic functions of time. The average number of species per m² decreased in all plots, in some cases as much as 50%. It was concluded that the diversity had decreased as a consequence of decreased species richness, decreased evenness and decreased pattern diversity. Many individual species distributions showed a pattern of nuclei surrounded with marginal belts. Differences in rate of change and persistence of spatial patterns between the plots, could be attributed to the differences in management history. These differences have disappeared as the succession has proceeded. The two exclosures have instead both conformed to the same floristic gradient, in turn based on a similar pattern of environmental gradients, primarily moisture. In the observed changes the emphasis was on the shifting importance of competing species populations, as some gained in importance at the expense of others. Competition has so far been a more important process in the vegetation development than immigration/extinction rates.Nomenclature follows Lid (1974) for vascular plants and Nyholm (1954–69) for mosses.This paper is the third part of a doctor's thesis from the Department of Plant Ecology, University of Lund. I wish to thank Prof. Nils Malmer for valuable advice and discussions, Mrs Mimmi Varga for drawing the figures and all my colleagues at the Department for their help and support.     

Vegetation,land use and management of the inner-dune zone in the North-Holland Dune Reserve,The Netherlands

The inner-dune zone between the villages of Bakkum and Egmond in the North-Holland Dune Reserve is one of the few remaining examples of the semi-natural boundary between the dunes and the polders. For several centuries this area has been used for various purposes, in particular for grazing. Large-scale excavating, especially after about 1900, has encroached upon the dunes. Vegetation and land-use history were studied and a provisional scheme on the vegetation development is presented. Management policies discussed are non-intervention, mowing in order to start natural processes such as rabbit grazing, and the continuation of traditional livestock grazing. Proposals for external management policies were made in order to maintain or enhance the original transition zone, based on comprehensive physical-geographical studies of the whole inner-dune zone.     

The significance of waterfowl grazing in the primary vegetation succession on embanked sandflats

In the annual stage of a primary plant succession on desalinating sandflats of the Lauwerszeepolder, N Netherlands, grazing by waterfowl is a dominant factor, as is illustrated by exclosures. It is concluded from population dynamical data, that both the removal of biomass and the predation of seeds sustain the annual species and retard succession to wards the next stage of a perennial grassland by several years. Yearly mowing in September slightly enhances succession by selectively affecting the dominant annual Salicornia species. The recurrence of high salinities in the upper soil layer of bare, c.q. grazed surfaces is expected to limit the invasion of grass species and other perennials.Nomenclature of plant taxa follows Heukels/van der Meijden (1983), Flora van Nederland, 20th ed. Wolters-Noordhoff, Groningen.This article would not have been written without the contributions of many students and colleagues, equally aware of the rare opportunity we had to study a genuine primary succession. I wish to thank them all. The help in the moving experiment of J. Franke, S. Nijdam, the late M. Robertus, H. Vrieling and J. D. D. Hofman is gratefully acknowledged. Thanks are due to the State Authority of the IJssellake Polders, for permitting access to and use of the experimental areas.     

Interactions between Limonium vulgare and Plantago maritima in the Plantagini-Limonietum on the Boschplaat,Terschelling, The Netherlands

Limonium vulgare and Plantago maritima are growing together over large areas and in varying proportions on the Boschplaat salt marsh. The question is how both species interact and which environmental factors play a role in those interactions.There are indications of different measures of niche differentiation in the Plantagini-Limonietum association related to differences in flooding frequency and substrate. Interactions between both species are described according to three models (related to de Wit's competition model). Mechanisms that may cause niche differentiation are: differences in growth rhythm, in salt tolerance, in rooting depth, in preference to types of substrate, in morphology, in mycorrhizae and diseases.On a small scale (spatial heterogeneity in the marsh and variation in subsequent seasons) Limonium can achieve a bigger portion in the vegetation by occupying open sites faster than Plantago. On a larger scale (the whole marsh, decades) the impossibility of Plantago to germinate and rejuvenate after sedimentation of silt and salinisation of the soil is the determining factor causing an overall decrease of this species (compared with Limonium) and increase of Limonium.The life cycle of both species, criteria for scale in space and time, and the question whether the Plantagini-Limonietum should be considered a syntaxon, are discussed.Nomenclature follows Heukels-van Ooststroom (1977).I like to thank Dr. P. Ketner and Dr. C. J. T. Spitters for their accompaniment and their ideas, so that I was aware of the wood (Boschplaat) for the trees. I also like to thank Drs. A. J. M. Roozen and Prof. Dr. E. van der Maarel for their discussions about this subject on many occasions. Finally I would like to thank Dr. W. G. Beeftink for his ideas and support and Ron Don, Wim van Hooff, Sjef Jansen, Alex Jonker, Annemiek Demon, Wim Heijligers, Marlie Becks and Frans Wilms for providing the basic concept (the Plantagini-Limonietum) and for their help.     

Ski slope vegetation at Snoqualmie Pass, Washington State, USA, and a comparison with ski slope vegetation in temperate coniferous forest zones

Ski slope vegetation at Snoqualmie Pass in Washington State, USA, was surveyed in order to identify community types and to compare it with vegetation development patterns in Japan. Ski slopes in Japan, most of which were constructed after 1960, underwent heavy land recontouring, while those at Snoqualmie Pass were constructed before 1950 with less modification. Three points apply to Japanese ski slope vegetation and differentiate these slopes from those at Snoqualmie Pass: (i) grasslands of introduced species are widespread and persistent; (ii) unvegetated patches are uncommon; and (iii) wetland vegetation has developed. These differences are mainly derived from the intensity of human impact, history of the slope and its scale: namely, ski slopes in Washington are older and larger than those in Japan. Ski slope vegetation in Washington was primarily differentiated by a soil moisture gradient. The large size of Washington ski slopes permitted the inclusion and development of wetland habitats, whereas most ski slopes in Japan are constructed on ridges and do not contain wetlands. Most introduced species in Japan are eliminated soon after seeding. In contrast, the long-term management of ski slopes decreased soil erosion and/or unvegetated patches in Washington and created relatively permanent grasslands composed of introduced species. Tsuga heterophylla and Abies amabilis were found established on the ski slopes in Washington, whereas in Japan the pioneer tree species are shade-intolerant broadleaved species. These differences may be a result of the different disturbance histories of ski slopes in the two countries. In addition, along with the conifers, early successional forbs such as Anaphalis margaritacea and Epilobium angustifolium are well established on Washington ski slopes. Results show that disturbances created by ski slope development greatly affect the vegetation, even on older, less heavily impacted ski slopes.     

A vegetation map of The Netherlands,based on the relationship between ecotopes and types of potential natural vegetation

Summary The method of mapping the vegetation on scale 1: 200,000 and the starting points in relation to the potential natural vegetation and ecotopes, are discussed.In view of the planological background of this study, some restrictions have been added to the concept of potential natural vegetation, concerning the period of development and the human influence.The relationship between soil, ground water and vegetation was studied, which resulted in the map of the potential natural vegetation.Each type of potential natural vegetation stands for a series of vegetation types on the same site. Seven main series, with a number of sub-series are distinguished. Within each vegetation series the plant communities have been spread over five groups, according to their structure and naturalness.Ecotopes and ecotope complexes are considered as landscape ecological units. A list of ecotopes was obtained by interpreting topographical maps and by inventory data.The actual vegetation was mapped by estimating the size of the ecotopes within the separate areas. It was expressed in a five figure code for the five groups from the vegetation and ecotopes is combined into the vegetation map of The Netherlands.Interpretation problems, some of them specific for The Netherlands, are discussed and some remarks are made on the necessity of further research.Contribution to the Symposium on Plant Species and Plant Communities, held at Nijmegen, 11–12 November 1976, on the occasion of the 60th birthday of Professor Victor Westhoff.Nomenclature follows Heukels-van Ooststroom, Flora van Nederland, 18e druk, 1975, Wolters-Noordhoff, Groningen; nomenclature of syntaxa follows Westhoff & den Held (1969)     

Studies on the succession of vegetation on some islands and sand banks in the nile near Khartoum,Sudan

Summary The main Nile near Khartoum contains a number of islands which differ in age, degree of soil stabilization, height in relation to flood-water mark and in the amount and composition of plant cover. These islands were studied with a view to understanding the sequence of colonization and succession.The initial stages occur on the newly exposed sand bars. A very thin vegetation cover of Tamarix nilotica occurs. The main habitat factor is soil mobility and the open soil texture.The successive stages are all characterized by a progressive increase of the silt content, an improvement of the water relations of the soil and greater soil stabilization. This improvement in soil conditions is associated with a distinct trend in the vegetation towards an increase in the number of species, the height of the tallest species and the amount of plant cover. The corresponding communities are: thin Glineto-Ambrosietum, Tamaricetum and Acacieto-Tamaricetum. The climax (or subclimax) occurs on the Nile banks and is represented by dense Acacietum.The role played by man in this succession is not clearly understood. The scanty evidence suggests that man's activities may be at least partly responsible for the succession of Tamaricetum by Acacieto-Tamaricetum.     

Effects of scale-dependent factors on herbaceous vegetation patterns in a wetland,northern Japan

Herbaceous vegetation was examined in an Otanoshike wetland in northern Japan to clarify the relationships between vegetation patterns and environmental factors with different scales. Alders (Alnus japonica) have recently invaded and might modify the herbaceous vegetation. In total, 150 50×50cm plots were established on the transitional areas between alder thickets and grassy marshland. Cover was measured for the vascular plant taxa, and canopy area, number of stumps, number of mounds, water depth, elevation difference, litter thickness, soil organic matter, and soil pH were measured in each plot. TWINSPAN cluster analysis classified four vegetation groups: (i), grasslands represented by Phragmites australis, Trientalis europaea, Lythrum salicaria, and Hosta rectifolia; (ii), Calamagrostis langsdorfii, and Polygonum thunbergii grasslands with Spiraea salicifolia; (iii), reed swamp dominated by Phragmites australis, and (iv), marshland dominated by Carex lyngbyei. Canonical correspondence analysis indicated that water depth primarily divided vegetation groups 1–2 and 3–4. Alder established in drier sites mostly by re-sprouting, and the canopy affected light and soil conditions on the ground surface. The second axis of canonical correspondence analysis was related to the canopy area and soil pH, and explained the vegetation differentiation between groups 1 and 2, and groups 3 and 4. In conclusion therefore, scale-dependent or hierarchical variables affected the vegetation patterns in different ways, that is, the herbaceous vegetation was first differentiated by water depth that was corresponding to alder establishment on a large scale, and subsequent light and soil conditions were second determinants on a small scale.     

Edge effects on vegetation and soils in a Virginia old-field

We investigated the effect of proximity to forest edge on plant community structure and ecosystem properties during succession, using field measurements of leaf area index (LAI), species composition, and soil carbon. Data were collected along four transects within a 14-year-old temperate successional field in north-central Virginia over the 2000 growing season. Additionally, the normalized difference vegetation index (NDVI) was calculated from LANDSAT 7 satellite data at a resolution of 30 m for the entire field. Results showed that relative frequencies of trees increased with proximity to forest edge suggesting a more advanced stage of succession in areas close to the adjacent secondary forest. Significant negative relationships were observed between distance from forest edge and both peak season LAI and NDVI. LAI and NDVI values within 60 m of the forest edge, however, were not significantly different from those values in the adjacent mature secondary forest, suggesting that some community level properties may take relatively short periods of time to reach undisturbed states. The presence of several key plant species, particularly Celastrus scandens (climbing bittersweet), exhibited a strong control on the spatial variability of LAI and potentially the aboveground net primary production. Soil carbon levels did not show a significant increase at sites close to the adjacent secondary forest (relative to an adjacent crop field), as seen with LAI and NDVI, suggesting no recovery of soil carbon in these systems after 14 years. This study points to the complexity of factors that influence spatial patterns of succession in old-fields and suggests that invasive species may play an important role in successional pathways and carbon cycling.     

Nutrient limitations and their implications on the effects of atmospheric deposition in coastal dunes; lime‐poor and lime‐rich sites in the Netherlands

1 A survey of plant and soil parameters was carried out in dry dune grasslands along the Dutch coast in the lime‐ and iron‐poor Wadden district and initially lime‐ and iron‐rich Renodunaal district, in order to detect differences in nutrient availability related to soil characteristics and potential sensitivity to atmospheric deposition of nitrogen.
2 Plant biomass and phosphorus pools in the shoot were higher in the Wadden district. The low foliar nitrogen concentrations and nitrogen/phosphorus ratios in the Wadden district suggested nitrogen‐limitation, while in the Renodunaal district there appeared to be a balanced supply of both nitrogen and phosphorus.
3 Soil pH, soil organic matter, soil nitrogen and phosphorus concentrations and total amounts were generally higher in the Renodunaal district. In both districts mineral phosphorus decreased with acidification and phosphorus oxalate (iron and aluminium bound) increased.
4 In the Wadden district iron is present primarily in iron–organic matter complexes, which leads to reversible binding of phosphorus. In the Renodunaal district large amounts of iron (hydr)oxides occur and at high pH may contribute to reversible phosphorus‐sorption, but at low pH this probably leads to immobilization of phosphorus.
5 While pools of soil phosphorus are low in the Wadden district, the phosphorus availability may be relatively high due to the comparatively loose nature of phosphorus‐sorption. As a result the area may be nitrogen‐limited and grass‐encroachment may thus have resulted from atmospheric deposition of nitrogen.
6 In the Renodunaal district, atmospheric deposition probably only accelerates grass‐encroachment, because deposition of acid and nitrogen increases the availability of both nitrogen and phosphorus and maintains the 'co‐limitation'.     

Energy Efficiency and Structural Change in the Netherlands, 1980–1995

International agreement has been reached to reduce greenhouse gas emissions worldwide. One important way of decoupling CO₂ emissions from economic growth is by introducing technical measures to improve energy efficiency. In this article, we assess the influence of developments in energy efficiency and economic structure on the total primary energy consumption in the Netherlands over the period 1980– 1995. We find a distinct decoupling of the economic growth and energy consumption of 1.5% per year in the 15-year analysis period. We measure (technical) changes in energy efficiency by changes in the energy consumption per physical unit of production or activity. The aggregate rate of (technical) energy-efficiency improvement was 1.4% per year over the period 1980–1995. The use of physical production indicators makes it possible to measure energy-efficiency developments without detailed surveys at a very low level of aggregation. When we look at economic structural changes over this period, we find that (i) no substantial shift took place at the level of the economic sectors that we distinguish; (ii) the most energy intensive subsectors grew much faster than the total economy; and (iii) at the subsector level, on average, a sizable decoupling of physical production and value added occurred. We conclude that structural changes, that is, changes in the composition of the economy, did not lead to a net decrease in the energy intensity of the Netherlands over the period 1980–1995.     

中国东北西部地区沙质荒漠化过程与植被动态关系的生态学研究：植被的排序

根据东北沙质荒漠化地质呼伦贝尔沙地,科尔沁沙地的72个沙地植物群落样地,276种植物的调查数据进行DCA排序,排序结果充分反映了沙地植被与地理因子,气候因子,地形因子,水文因子,干扰因子的关系,DCA排序的第1轴反映沙地植被分布的水分梯度;第2轴反映土壤梯度;第4轴主要反映热量梯度;DCA的1、4轴排序图和1,2,4轴排序图,反映沙地植被与沙质荒漠化的生态规律,采用定性因子评价赋值与定量因子相结合的方法,经双重筛选逐步回归分析,建立了沙地植被DCA排序坐标与地理,气候,地形,水文,干扰等诸多生态因子的线性回归模型。