Fluctuations in a coastal dune grassland due to fluctuations in rainfall: Experimental evidence

Experiments with rainfall on a dune grassland near Oostvoorne, the Netherlands with Festuco-Galietum as the main syntaxon are described. Both increase in rain through additional watering and decrease in rain through catchment are presented to plots belonging to the xerosere and the mesosere. Clear changes in the floristic composition are the result, even after only two years. Typical Festuco-Galietum species are promoted by high rainfall, species of open habitat such as Corynephorus canescens are promoted by drought. The results are discussed against the background of long term permanent plot observations in the area and a relation with rabbit grazing intensity is supposed.Nomenclature of vascular plants follows Heukels-van Ooststroom (1975) Flora van Nederland 18 ed., Woiters-Noordhoff, Groningen; Nomenclature of syntaxa follows Westhoff & den Held (1969). Plantengemeenschappen in Nederland, Thieme, Zutphen.Field work in 1978 was carried out with Frans Bongers and Marc de Lyon, in 1979 with Marc de Lyon, Picter Meeuwissen and Guiljo van Nuland, all then MSc. students at the Division of Geobotany. Their help and the advice of Dr. Peter van der Aart, Institute of Ecological Research, Oostvoorne, are acknowledged.     

The effects of dissolved macro-nutrients on the herbaceous vegetation around dune pools

In Dutch coastal dunes the relation was studied between the composition of the vegetation on the banks of pools and the nutrients supplied by the groundwater. Areas infiltrated with eutrophic river water were compared with non-affected areas. The floristic composition was interpreted according to classification systems of ruderal, nitrophilous tall hemicryptophytes and the conservation parameters richness and rarity. Of the three main macro-nutrients (orthophosphate, nitrate and potassium), the orthophosphate load shows the highest correlation with the vegetation variables. It is suggested that a reduction of orthophosphate during pre-purification of the infiltration water would reduce the dominance of the ruderals to the extent that the original dune slack species could return locally.Nomenclature of species follows Arnolds & Van der Meijden (1975).We thank G. J. de Bruyn, W. H. van Dobben, W. T. de Groot, W. J. ter Keurs, F. van der Meulen, W. H. van der Molen, M. A. W. Noordervliet, and I. S. Zonneveld for their comments on the study; J. Braak, J. Brakkee and R. van Nood for their assistance with the chemical analyses; A. R. Kaal for the translation, and G. P. G. Hock for the drawing of the figures.     

Spatial and temporal variation in the vegetation of dune slacks in relation to the ground water régime

Summary The aim of the study presented here is to determine the relation between the vegetation and the ground water régime in the dune slacks on the island of Voorne. A distinction is made between the spatial and the temporal variation in the vegetation. On the basis of data from a transect the relation between the spatial variation and the water-table is determined. With respect to the temporal variation data recorded on permanent plots were used. Both the variation in space and that in time proved to be related to differences in the ground water régime. The supposition is put forward that the amount of variation in time depends on the height of site relative to the water-table.Contribution to the Symposium of the Working Group on Succession Research on Permanent Plots held at Yerseke, the Netherlands, October 1975.Nomenclature of phanerogams follows Heukels & van Ooststroom 1975, Flora van Nederland, 18e druk Wolters-Noordhoff, Groningen; that of bryophytes Margadant 1959, Mossentabel, 3e druk, Amsterdam.The author is very much indebted to Mr. P.A. Bakker, Society for the Preservation of Nature reserves in the Netherlands, for the use of the 1966 relevé of site 46, and to Mr. W. Smant for his assistance in field work and the preparation of the data.     

Size hierarchy and mineral status of Rhinanthus angustifolius populations under different grassland management regimes

The size hierarchy and mineral concentrations of adult plants in flower and their seed crop have been investigated in three populations of Rhinanthus angustifolius under a grassland management regime consisting of haying without fertilization. The effect of this type of management on the nutrient budgets of soil or Rhinanthus plants was shown to be small initially, i.e. up to the twelfth year of changed management. Although the mean biomass of plants significantly diminished by 30% in this period, their nutrient concentration was not strongly affected. Small plants consistently had lower concentrations of N and Na than large plants. Prolongation of this management regime beyond twelve years let to a general reduction in the phosphorus concentration of the plants. An observed increase in the concentrations of K, Mn, and Cu, especially in small plants, is tentatively ascribed to an imbalance in mineral nutrition.Nomenclature of vascular plants follows Van der Meijden et al. (1983). Flora van Nederland.     

Some considerations of and perspectives in coastal ecology

Ecophysiological studies of plant species in wet saline habitats have been one-sided in the past, emphasizing sodium chloride only. More emphasis should be given to anaerobiosis, to mechanical factors, to the effects of a surplus of other nutrients in seawater and coastal ecosystems and their metabolic impact as well as to aspects of nutrient (re)allocation. Research should incorporate theoretical models on plant strategies maximizing yield, minimizing risk. At the population level, a new operational parameter for ecological optima and ecological stress is presented considering normal and skewed distributions of plant parameters in relation to growth and the concentration of mineral nutrients.Some considerations on the ecological effects of phenotypic plasticity and genetic variability are given by emphasizing the aspects of diaspores.Future investigations of coastal vegetation should consider the interaction of populations within plant communities, including the regulating effects of animals.Nomenclature of vascular plants follows H. Heukels & R. van der Meijden, 1983. Flora van Nederland, 20th ed. Wolters-Noordhoff, Groningen; that of plant communities V. Westhoff & A. J. den Held, 1969. Plantengemeenschappen in Nederland. Thieme & Cie, Zutphen.     

Vegetation development in a former orchard under different treatments: A preliminary report

Summary Vegetation development in a former orchard on loamy moist soil after removal of the fruit trees under different treatment is described with the help of multivariate methods. Data on 20 plots over 9 years are treated with a simple classification program and a PCA ordination. Two main groups of plots are involved: one in a relatively moist part of the area where the trees were removed in 1970, one in a slightly drier area where the trees had been removed earlier. The first group is characterized by forbs such as Urtica dioica and Lamium maculatum, the second by grasses, mainly Arrhenatherum elatius.Under a mowing regime the plots of the first group change in the direction of the second group. Under continued fertilization the Urtica vegetation can be maintained. Establishment and/or rapid spread of Urtica can be brought about in Arrhenatherum dominated sites by (re-) introduction of nitrogen fertiliser.Not mowing leads to a dense Urtica-Lamium stand in which Lamium maculatum finds an optimum some years after mowing is stopped.The methods of summarizing the site-time data are discussed and some new ones announced for further research.Nomenclature follows Heukels-van Ooststroom, Flora van Nederland, 18e druk, Wolters-Noordhoff, Groningen, 1975.Vegetation and soil analyses were mainly carried out by students in the framework of their study for the degree of doctorandus in biology: Ineke Opbroek, Ferdinand Baggerman (1970), Carla Veldhuis (1971–1972), the late Doeke Dijken (1971–1973), Jos Rijpert (1972), the Rev. Piet van der Aart, Jo Louppen (1973), Frank Verhoeven (1974), Maurits Heine (1975), Ad Peters (1976), Huub Muyres, Louis Fliervoet (1976, 1977), Emilia Peters (1977), Niek Joanknecht, Thijs Lavrijsen (1977, 1978).     

Water status,drought responses,and growth of <Emphasis Type="Italic">Prosopis flexuosa</Emphasis> trees with different access to the water table in a warm South American desert

Prosopis flexuosa trees dominate woodlands in the Central Monte Desert (Mendoza, Argentina), with <200 mm rainfall, exploiting the water table recharged by Andean rivers, and also growing in dunes with no access to the water table. Prosopis woodlands were extensively logged during development of the agricultural oasis, and surface and groundwater irrigation could lower the depth of the water table in the future. We evaluated tree populations with decreasing access to the water table: valley adult trees, valley saplings, and dune adult trees, in order to assess their ecophysiological response to water table accessibility. High and seasonally stable pre-dawn leaf water potentials (−2.2 ± 0.2 to −1.2 ± 0.07 MPa) indicated that valley adults utilize larger and more stable water reservoirs than valley saplings and dune adults (−3.8 ± 0.3 to −1.3 ± 0.07 MPa), with higher midday leaf conductance to water vapor (valley adults ~250; dune adults <60 mmol m⁻² s⁻¹), potentially higher CO₂ uptake, and increased radial growth rate (valley adults 4.1 ± 0.07; dune adults 2.9 ± 0.02 mm year⁻¹). Trees with poor access to the water table exhibited drought tolerance responses such as midday stomata closure, leaflet closure, and osmotic adjustment. Stomata density decreased in response to drought when leaf expansion was restricted. The combination of phreatophytism and drought tolerance would enlarge P. flexuosa habitats and buffer populations against changes in rainfall dynamics and water table depth.     

The reaction of some sand-dune plant species to experimentally imposed environmental change: a reductionist approach to stability

Coastal dunes in western Europe have long been under grazing and recreational pressure, but the vegetative cover appears to be remarkably resilient. In a series of experiments on Ynyslas Dunes, Cardigan Bay (Wales, U.K.), trampling, protection, fertiliser, and herbicide treatments were monitored using non-destructive and destructive sampling methods. This paper discusses the behaviour of 12 important species after these treatments on three major dune habitats: Agrostis tenuis, Ammophila arenaria, Bellis perennis, Festuca rubra, Galium verum, Leontodon taraxacoides, Ononis repens, Plantago lanceolata, Poa subcaerulea, Thymus praecox, Homalothecium lutescens, Tortula ruraliformis.Although stability is usually thought of as a feature of the community, a reductionist approach suggests that grasses react quickly but temporarily, dicotyledonous forbs often show slower reaction times, each species reacting in its own way to changes in the environment. A tentative classification of our study species is possible.Nomenclature follows Smith (1978), for bryophytes and Clapam, Tutin & Warburg (1962), for angiosperms.     

Multi-species pattern analysis of Mediterranean pastures in three stages of ecological succession

Multi-species patterns were analyzed in three stages of ecological succession in oligotrophic pastures in Central Spain using a new method of analysis. The method is based on plotting the species coordinates of principal components analysis undertaken at different block sizes. It allows the detection of any heterogeneity present and reveals both microtopographical and macrotopographical trends in species patterns.Nomenclature follows Tutin et al., 1964–1980, Flora Europaea.     

Shoot biomass and species richness in relation to some environmental factors in a coastal dune area in The Netherlands

Shoot biomass, species richness and selected environmental factors were studied at Meijendel, a coastal dune area in The Netherlands. The relationship between species richness and shoot biomass of the stand could be best described by a unimodal curve, with the peak at 300 g m^–2. Measures of species richness were positively correlated with soil water, nitrogen and phosphorus.Shoot biomass showed a positive correlation with soil water, nitrogen and humus at low levels of shoot biomass.Nomenclature of phanerogams and plant community types follows Heukels & van Ooststroom (1973), and Westhoff & den Held (1969) respectively.Publication of the Meijendel comité new series: 76.     

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.     

Aspects of population dynamics in Halimione portulacoides communities

Summary Studies on sample plots inHalimione portulacoides communities show that environmental disturbances, either natural or induced by man, start a sequence of partly overlapping density maxima inSuaeda maritima, Aster tripolium andPuccinellia maritima successively, before the originalHalimione community totally recovers. When succession time before recovering is long enough, there are tendencies in redundancy of this sequence stressing the unilinear character of the succession. Minor environmental impacts induce a longer time-lag period of theSuaeda density maximum, suggesting threshold values of these impacts for the species to maintain minimal population densities or to become locally extinct. This sequence of interim species starting after an environmental disturbance, suggests also a gradient character in various biological attributes, for instance in life-time, propagation, nutrient and genetic plasticity strategies. The mechanism described can therefore be interpreted as a complex of mostly well-adapted and well-integrated inherent species strategies capable of absorbing environmental shocks. It is suggested that in the salt-marsh ccosystem the pattern of spatial variation in densities and that of temporal variation in fluctuations of the three species populations under natural conditions reflect corresponding patterns of environmental disturbances in the vegetation taking into account a timelag associated with the magnitude of the impact concerned.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-Noodhoff, Groningen.The authors are greatly indebted to Dr K.F. Vaas (Yerseke) for reviewing the English text.Communication Nr. 160.     

Interspecific relationships of the glacial relics Swertia perennis L. and Pedicularis sceptrum-carolinum L. in a Rumanian fen

In a eutrophic peat bog impregnated with mineral springs in the Carpathians, where the glacial relics Swertia perennis and Pedicularis sceptrum-carolinum have earlier been shown to occupy different microhabitats, an analysis of the relationships of these species with accompanying species has been made on the basis of numerous quadrats and a correlation method. The closest floristic associates of the two species are quite different, and the results of this analysis are broadly in agreement with subjective assignments made earlier by phytosociologists.Nomenclature follows T. G. Tutin & V. H. Heywood (eds.), Flora Europaea, Vols. I–V. Cambridge Univ. Press, Cambridge, 1964, 1968, 1972, 1976, 1980 for Cormophyta and R. van der Wijk (ed.), Index Muscorum, Vols. I, II. Utrecht, 1959, 1962 for Bryophyta.     

Micro-patterns in grassland vegetation created and sustained by sheep-grazing

An initially uniform Holcus lanatus-dominated sward came partly under hay-making and partly under sheep-grazing. Preferential grazing by sheep resulted in grazing at different intensities giving rise to a macro-pattern of various plant communities. Besides this macro-pattern a micro-pattern developed in the grazed area, which was absent under hay-making. In the micro-pattern short, heavily grazed areas alternated with taller, lightly grazed patches, both having the same species composition. The heavily grazed area was characterized by equal amounts of monocots and dicots. The lightly grazed patches were dominated by Agrostis tenuis, and had a large amount of litter which probably causes the absence of mosses. The protein percentage of green material is higher in the heavily grazed areas than in the lightly grazed patches.Sequential charting indicated that the micro-pattern was more or less stable. An interaction between the vegetation micro-pattern and grazing patterns is suggested. Heavy grazing results in forage with a high protein content and hence attracts animals. Light grazing results in forage with a relatively low protein content, animals avoid the area and litter accumulates.Nomenclature follows Heukels & van Ooststroom (1977) Flora van Nederland.Mrs J. O'Brien corrected the English text     

Successive mapping of dune slack vegetation

Summary The succession of the dune slack vegetation around an artificial lake was studied by means of successive mapping. A succession scheme was prepared on the basis of comparison of the maps made in various years. This scheme was discussed in terms of systems theory, with reference to a kinematic graph prepared on the basis of the transformations. The succession derived from the vegetation maps was compared with that derived from the permanent plots; the results of this comparison indicated that only a small proportion of the total number of succession lines could be detected with the aid of the permanent plots. In addition, the information that could be derived from the permanent plot analysis depended on the degree of change of the vegetation in the area; more information could be derived from drier areas which showed the least degree of change. The best method for the investigation of the succession proved to be a combination of a restricted number of permanent plots and successive mapping. Nomenclature of species follows Heukels-van Ooststroom Flora van Nederland, 16 ed. 1970. R.I.N.-Comm. nr. 107.     

Rabbit grazing and species diversity in a dune area

Summary The relation between density of rabbit populations and plant species diversity is discussed. In the dunes of the Dutch Frisian Island of Schiermonnikoog, the rabbit grazing pressure has been quantified on the basis of traces of recent grazing activity. Moderate grazing turned out to bring about maximal species richness. Current changes in grazing pressure (either decrease or increase) can result in plant species diversity that is lower than in other places with equal but stabilized grazing pressure.Nomenclature follows Heukels-van Ooststroom, Flora van Nederland, 18e druk, 1975.The work has been done by Mrs Ada Zeevalking-van Y peren and Mr Hein Zeevalking as part of their study for a doctoral degree.     

Air pollution as a possible cause for the decline of some phanerogamic species in The Netherlands

In The Netherlands the decline of some phanerogamic species cannot be readily explained from obvious factors such as lowering of the groundwater table, eutrophication or land reclamation. For a number of species the hypothesis was tested that the decline is partly due to air pollution. A two-factor model was made in which decline is accounted for by (a) habitat destruction assessed from topographic maps and (b) air pollution measured as the SO₂ 95-percentile over the winter period 1978/1979. Effects of both factors were assumed to follow a sigmoid dose-effect curve. For a number of species decline proved to be significantly correlated with air pollution. These are notably species from the syntaxon Violion caninae. A comparison was made with results obtained for epiphytic lichens. It appears that for some phanerogamic species sensitivity is about the same as for moderately sensitive lichens.Nomenclature follows Heukels & van der Meijden (1983).Thanks are due to the Rijksherbarium, for providing some of their unpublished data; and to Ada Groeneveld, for technical assistance.     

山东水龙骨科植物孢粉学研究及其在分类上的意义

基于Flora of China采用国际石松类和蕨类植物新分类系统,其水龙骨科(Polypodiaceae)不仅涵盖了秦仁昌分类系统中的水龙骨科(Polypodiaceae) 20多属,而且还包括槲蕨科(Drynariaceae)、鹿角蕨科(Platyceriaceae)等独立的科。水龙骨科(Polypodiaceae)新分类系统种类较多、分类复杂,需要对其合理性和孢粉学在新分类系统中的意义进行新的探讨。该文首次以新分类系统为依据,采用扫描电镜对山东分布的水龙骨科(Polypodiaceae)植物孢子形态及其周壁纹饰亚显微结构进行了系统地观察。结果表明:该科孢子形态为圆肾形,左右对称,具周壁,其周壁纹饰在种内稳定,在亚科、属及种间区别显著。孢粉学研究结果,支持槲蕨亚科、鹿角蕨亚科分别列于水龙骨科下的两个亚科;基于山东假瘤蕨孢子周壁纹饰,并结合叶片形态特征,山东假瘤蕨(Phymatopteris shandongensis J.X.Li et C.Y.Wang)应为一个独立的新种,不宜并入金鸡脚假瘤蕨[Phymatopteris hastata (Thunb.) Pic.Serm.]。依据《国际植物命名法规》和Flora of China分类系统,山东假瘤蕨(Phymatopteris shandongensis J.X.Li et C.Y.Wang)新组合为山东假瘤蕨[Selliguea shandongensis (J.X.Li et C.Y.Wang) J.X.LiX.J.Li,comb.nov.]。该研究结果不仅首次为水龙骨科植物孢粉学积累了新资料,而且也为其新分类系统提供了孢粉学的科学依据,与DNA分类系统相吻合,证明其新分类系统的合理性和科学性,在构建水龙骨科及其亚科的自然分类系统中具有重要意义。     

Life history and population dynamics of Atriplex triangularis

The dynamics of a population of the stoloniferous Potentilla anserina was studied during four years. The mortality of established ramets followed a log-linear pattern superimposed on a seasonal fluctuation with an increased mortality during the period of vegetative spread. The major external mortality factors were unfavourable weather conditions such as drought and freezing. The turnover rate of the population was slow (31.6 yr). Vegetative spread dominated the recruitment to the population. Seedlings occurred but were exposed to a much higher mortality than vegetative daughter ramets. The juvenile period of seedlings is at least 4 yr in the field. This contrast to vegetative daughter ramets which are potentially flowering and stoloniferous the year after they are established, even if age affects both reproduction and vegetative spread. The density of most of the population is below the level where intraspecific regulative effects operate. However, in very dense stands both stolon and flower production were negatively density dependent.Nomenclature follows Lid (1974), Norsk og Svensk Flora, 2nd ed. Oslo.I am indebted to Lenn Jerling, Peter Torstensson, Anders Telenius and Gudrún Jónsdóttir for their help, and inspiring criticism throughout this study.     

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)