Jan P. Bakker

Constraints in the restoration of ecological diversity in grassland and heathland communities

Species-rich grassland and heathland communities still occur in low-intensity farming systems in many European countries. Gradually, such systems have either been abandoned or more intensively exploited, with a subsequent decrease in species numbers. Until recently, it was thought that restoration of these communities would be straightforward. However, abiotic constraints (with respect to eutrophication and acidification) have hampered restoration more than previously thought. Moreover, very recent research has revealed that biotic constraints can also be important: many plant species are not present in the soil seed bank and their dispersal is limited in the present fragmented landscape.

Nature Management by Grazing and Cutting

One: Introduction.- The need for nature management.- I Introduction.- II Research approach and methods.- Two: Land Use Dynamics.- From agricultural practices to nature management.- III The Drenthe Plateau.- IV The Schiermonnikoog salt marsh.- Three: Vegetation Dynamics.- Restoration management creates species-richer plant communities: three cases.- V The heathland and reclaimed grassland vegetation.- VI The valley grassland vegetation.- VII The salt marsh vegetation.- Four: Ecosystem Dynamics.- Causes of observed changes: mineral removal/increasing soil salinity or structure of the sward?.- VIII Grazing.- IX Cutting regimes.- X Changing soil conditions and stand structure as causes of floristic changes: conclusions and synthesis.- Five: Population Dynamics.- Where do the new species come from?.- XI Seed dispersal and seed bank.- XII Re-introduction of seeds.- Six: Management.- Evaluation of the obtained knowledge.- XIII Management.- Summary.- References.

An improved method for seed-bank analysis : Seedling emergence after removing the soil by sieving

1. The seedling emergence method for assessing the size of the seed bank is improved by washing soil samples on a fine sieve and spreading the thus concentrated samples in a 3-5 mm thick layer on sterilized potting compost. 2. The method largely increases the number of seedlings that emerge as compared to unconcentrated samples. Hand-sorting afterwards shows that the germination rates vary between 81 and 100% of the viable seeds present. 3. Ninety-five per cent of the seedlings will emerge within 6 weeks using this method. 4. The method greatly reduces the greenhouse space needed and enables examination of large sample volumes.

Relationship between soil chemical factors and grassland diversity

Many studies carried out during these last few years have focused on the factors influencing plant diversity in species-rich grasslands. This is due to the fact that these ecosystems, among the most diversified in temperate climates, are extremely threatened; in some areas, they have almost disappeared. The re-establishment of these habitats implies to know the living conditions of the associations to be recreated. Very often, the typical species of these communities have become so rarefied that the seed bank or the seed rain are not sufficient to recreate the plant community. Most of the time, to achieve the restoration of these communities, they have to be totally recreated by sowing. For the restoration or the maintenance of the community, the soil chemical characteristics have also to be appropriate or if not modified. This research tends to establish a relation between some soil chemical factors and the plant diversity of a great number of stations. This research has illuminated the relationship between soil extractable phosphorus and potassium and plant diversity. Over 5 mg of phosphorus per 100 g of dry soil (acetate + EDTA extraction), no station containing more than 20 species per 100 m2 has been found. The highest number of species is found below the optimum content of the soil for plant nutrition (5–8 mg P/100 g). Concerning the potassium, the highest number of species is found at 20 mg/100, a value corresponcing to an optimum content of the soil for plant nutrition. High potassium contents, in opposition to phosphorus contents, are thus compatible with high values of diversity. Other factors (i.e. pH, organic matter, total nitrogen and calcium) do not show so clearly a relation with plant diversity. Excess of N–NO3 is known for its negative effect on the diversity of plant communities. In these environments, apart from the atmospheric deposits which can be important in some areas, N–NO3 is derived mainly from the symbiotic fixation of atmospheric nitrogen by legumes as well as from the mineralization of the organic matter of the soil. It is possible that, when in small quantities, the available soil phosphorus could be a limiting factor of the N–NO3 supply by these two sources. In this hypothesis, nitrogen would remain the main element limitating plant diversity but its availability would be controlled by phosphorus.

Why do we need permanent plots in the study of long-term vegetation dynamics?

This paper presents a survey of vegetation studies on permanent plots, with an emphasis on the long-term char- acter of these studies. It makes remarks on the external and internal causes of succession, pays attention to the pathways of succession, links up permanent plots studies and chrono- sequences, discusses internal causes and mechanisms of suc- cession, and finally mentions the significance of long-term vegetation dynamics for nature conservation.

Vegetation succession and herbivory in a salt marsh: changes induced by sea level rise and silt deposition along an elevational gradient

1 The relationships between soil development, vertical vegetation zonation, vegetation succession and herbivory by Brent geese, Branta bernicla, were studied in a coastal salt marsh. We were able to analyse up to 100 years of salt marsh development by comparing sites where vegetation succession had progressed for-various periods of time. These data were related to a continuous daily record of high water levels measured since 1824. 2 Most elevational variation in edaphic conditions (and therefore vertical vegetation zonation) could be attributed to variation in height of the sandy subsoil, as rapid dune formation occurs on the beaches early in succession. In the intermediate part of this elevational gradient, the maximum annual increase of 1.2 mm of silt corresponded to an annual increase of 5.6 g N m(-2) in the topsoil (0-50 cm). The average sea level rise in this area over the last 170 years was 0.63 mm year(-1). A sedimentation model suggests that this has had strong effects on sedimentation and the annual inundation frequency in the mid-part of the elevational gradient, thus affecting vegetation zonation on the salt marsh. For the major part of the investigated transects, sea level rise has probably speeded up succession due to an increased rate of sedimentation. 3 The occurrence and dominance of all plant species were recorded in 3927 plots, and and for the 11 most common species response surfaces were calculated for their dependence on elevation and transect age. Most plant species were clearly separated along these axes. Most halophytic species, which were preferred by the geese, occurred early in succession and low on the gradient, where we observed the highest densities ge quality of Festuca I rubra increased towards the lower salt marsh. Other preferred forage species (Puccinellia maritima and Plantago maritima) were gradually displaced during succession by the tall grass Elymus athericus, especially in the mid-and upper salt marsh. Few geese grazed in areas where Elymus was dominant. 4 Herbivores first increased in numbers but then declined along a gradient of primary productivity. We propose that declining forage quality (due to changing vegetation composition during succession) is a better explanation for this pattern than the classic explanation of predator control of herbivores at high levels of primary productivity. This quality threshold hypothesis, as an alternative explanation of the exploitation ecosystem hypothesis, is expected to hold especially where smaller (quality-sensitive) herbivores such as geese are present. 5 Grazing by cattle in a 200-year-old part of the salt marsh led to the disappearance of Elymus athericus, to establishment of early successional halophytes and a return of Brent geese. Crazing by a larger herbivore therefore facilitated conditions for smaller herbivores by preventing the dominance of plant species that were good light competitors, and thus improved forage quality. Populations of these small herbivores could then become regulated by predators, although none was present at our site.

Grazing and Conservation Management

Theoretical background the development of grassland communities in north-western Europe evaluation - changes in plant species richness changes in plant communities structural diversity and boundary effects effects on the fauna effects on soil characteristics (nutrients, decomposition, accumulation of litter) changes in Dutch landscapes in relation to mangement options limiting factors - animal performance and carrying capacity grazing in relation to other environmental factors (hydrology, acid rain, plant geography, time) Conclusion and perspectives.

Self‐Organization and Vegetation Collapse in Salt Marsh Ecosystems

Complexity theory predicts that local feedback processes may strongly affect the organization of ecosystems on larger spatial scales. Whether complexity leads to increased resilience and stability or to increased vulnerability and criticality remains one of the dominant questions in ecology. We present a combined theoretical and empirical study of complex dynamics in mineralogenic salt marsh ecosystems that emerge from a positive feedback between clay accumulation and plant growth. Positive feedback induces self‐organizing within the ecosystem, which buffers for the strong physical gradient that characterizes the marine‐terrestrial boundary, and improves plant growth along the gradient. However, as a consequence of these self‐organizing properties, salt marshes approach a critical state as the edge of the salt marsh and the adjacent intertidal flat becomes increasingly steep and vulnerable to wave attack. Disturbance caused, for instance, by a storm may induce a cascade of vegetation collapse and severe erosion on the cliff edge, leading to salt marsh destruction. Our study shows that on short timescales, self‐organization improves the functioning of salt marsh ecosystems. On long timescales, however, self‐organization may lead to destruction of salt marsh vegetation.

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.

Soil seed bank composition along a gradient from dry alvar grassland to Juniperus shrubland

Soil seed bank composition along a gradient from dry alvar grassland to Juniperus shrubland.