Jelte van Andel

Effects of fire and herbivory on the stability of savanna ecosystems

Savanna ecosystems are characterized by the co-occurrence of trees and grass- es. In this paper, we argue that the balance between trees and grasses is, to a large extent, determined by the indirect interactive effects of herbivory and fire. These effects are based on the positive feedback between fuel load (grass biomass) and fire intensity. An increase in the level of grazing leads to reduced fuel load, which makes fire less intense and, thus, less damaging to trees and, consequently, results in an increase in woody vegetation. The system then switches from a state with trees and grasses to a state with solely trees. Similarly, browsers may enhance the effect of fire on trees because they reduce woody biomass, thus indirectly stimulating grass growth. This consequent increase in fuel load results in more intense fire and increased decline of biomass. The system then switches from a state with solely trees to a state with trees and grasses. We maintain that the interaction between fire and herbivory provides a mechanistic explanation for observed discontinuous changes in woody and grass biomass. This is an alternative for the soil degradation mechanism, in which there is a positive feedback between the amount of grass biomass and the amount of water that infiltrates into the soil. The soil degradation mechanism predicts no discontinuous chang- es, such as bush encroachment, on sandy soils. Such changes, however, are frequently ob- served. Therefore, the interactive effects of fire and herbivory provide a more plausible explanation for the occurrence of discontinuous changes in savanna ecosystems.

Pollen and gene flow in fragmented habitats

. Habitat fragmentation affects both plants and pollinators. Habitat fragmentation leads to changes in species richness, population number and size, density, and shape, thus to changes in the spatial arrangement of flowers. These changes influence the amount of food for flower-visiting insects and the quantity and quality of pollinations. Seed set in small populations is often reduced and genetic variation is expected but not always found to be low. The majority of studies show that low flower densities have reduced pollination success and higher inbreeding. Density effects are stronger than size effects. Most studies concluded that species richness in flower-visiting insects is directly related to richness in plant species. However, the consequences of low insect species richness for pollination are not always clear, depending on the studied pollinator-plant relationship. The effects of the presence of simultaneously flowering species are highly dependent on the circumstances and may range from competition to facilitation. Other flowering plant species may play a role as stepping stones or corridor in the connection between populations. In the absence of stepping stones even short distances between populations act as strong barriers for gene flow. We illustrate the present review paper with own data collected for three plant species, rare in The Netherlands: Phyteuma spicatum ssp. nigrum (Campanulaceae), Salvia pratensis (Labiatae) and Scabiosa columbaria (Dipsacaceae). The species differ in their breeding systems and in the assemblage of visitor species. Data are shown on the effects of population size on species richness with consequences for seed set. Effects of flower density and isolation on pollen exchange are given. Since plant reproduction depends on the behaviour of individual insects and not on the overall behaviour of the species, the examples all point to individual insects and extrapolate to effects at the species level.

Spatial Heterogeneity and Irreversible Vegetation Change in Semiarid Grazing Systems

Recent theoretical studies have shown that spatial redistribution of surface water may explain the occurrence of patterns of alternating vegetated and degraded patches in semiarid grasslands. These results implied, however, that spatial redistribution processes cannot explain the collapse of production on coarser scales observed in these systems. We present a spatially explicit vegetation model to investigate possible mechanisms explaining irreversible vegetation collapse on coarse spatial scales. The model results indicate that the dynamics of vegetation on coarse scales are determined by the interaction of two spatial feedback processes. Loss of plant cover in a certain area results in increased availability of water in remaining vegetated patches through run‐on of surface water, promoting within‐patch plant production. Hence, spatial redistribution of surface water creates negative feedback between reduced plant cover and increased plant growth in remaining vegetation. Reduced plant cover, however, results in focusing of herbivore grazing in the remaining vegetation. Hence, redistribution of herbivores creates positive feedback between reduced plant cover and increased losses due to grazing in remaining vegetated patches, leading to collapse of the entire vegetation. This may explain irreversible vegetation shifts in semiarid grasslands on coarse spatial scales.

Responses of fen plant species to groundwater level and light intensity

Characteristic species of sedge-moss fen communities occur in constantly wet, nutrient-poor sites with a high penetration of light through the vegetation canopy. We studied the effects of water table depth and differences in light intensity on the performance of fen species. Three fen species (Carex curta, Viola palustris, Hydrocotyle vulgaris) and one species with a wide range of occurrence (Poa trivialis) were grown for 10 weeks in a sedge-moss peat substrate at 4 different water levels and 3 light intensities. In all species differences in light availability had a larger effect on biomass production than differences in water level. Under a light availability reduced to only 10% the root weight ratio of all the species decreased while leaf weight ratio increased. The biomass allocation ratios were hardly affected by differences in water level. For Viol a and Hydrocotyle an interaction between the two factors was observed. Poa did not show particular differences compared to the other species. We discuss the results in the context of the establishment of fen species in riparian vegetation. It is suggested that the occurrence of fen species in the landscape is directly related to the availability of light, whereas the relationship between fen species occurrence and hydrological conditions seems to be an indirect one.

Intraspecific variability in the context of ecological restoration projects

Abstract Population differentiation within species is a common phenomenon. The question raised in this review is whether such variability should be recognized while making a choice for populations as sources for the (re)introduction of species into former or novel environments. To estimate the ecological importance of population differentiation in the context of restoration projects, I reviewed literature in which the results from (reciprocal) sowing and transplant experiments are reported. This survey reveals that survival of the transplants (one of the earliest recognizable fitness components) is almost entirely site-dependent, while the fitness of surviving plants (measured in terms of growth and fecundity) is frequently higher for native plant populations as compared to alien populations of a species. Phenological timing is sensitive to selection and rather difficult to recover in an adequate way. As far as fitness reduction is concerned, it can be stated that this is a minor effect in comparison to the complete lack of a species from the restoration site; however, the credits and debits of quantity vs. quality of the members of a population cannot yet be estimated scientifically. Comparative experiments in restoration projects would help in solving this problem. The suitability of environmental conditions for the species and the accessibility of the dispersal units to the restoration site are actually thought to be the major constraints.

Ecological communities: conceptual problems and definitions

Abstract Three major problems with respect to ecological communities are tackled. The first is the problem of ambiguity, i.e. the fact that the term “community” is being used for various kinds of objects at different levels of organisation. We argue that this problem can be resolved by restricting use of the term “community” to sets of co-occurring organisms belonging to a single taxonomic phylum or class (plants, birds, insects, etc.) and by using the term “biocoenosis” for sets of organisms belonging to multiple phyla or classes and comprising the biotic components of ecosystems. We also argue that interaction between organisms is neither a necessary nor a sufficient condition for community membership, or, in other words, that communities may consist of both interacting and non-interacting organisms. The second and third problems are the boundary problem and the problem of heterogeneity, i.e. the fact that communities, as currently defined, most often do not have discrete boundaries and are quite heterogeneous with respect to species composition. We argue that both problems result from the fact that communities are seen as groups of co-occurring populations of species, whereas actually populations of different species rarely if ever co-occur in exactly the same area. These problems can be resolved by redefining communities as particular sets of individuals occurring in the intersection of the areas occupied by different populations of species. In the final section we defend our definition against some potential objections, viz. (1) that it would lead to an excessive number of small communities, especially in species-rich situations, and (2) that, because population boundaries may change all the time, such communities would be very unstable.

Two approaches towards the relationship between plant species diversity and ecosystem functioning

. The main question to be dealt with in the papers published in this Special Feature is to which extent plant species richness can be applied as a parameter in restoration projects to qualify the ecosystems state. Before considering this problem, it should be recognized that this approach illuminates only one side of the coin; the other side is touched by the opposite question, asking which plant species are essential components of an ecosystem. These two approaches towards the relationship between species richness and ecosystem functioning are not mutually exclusive, but should not be confused either. In view of ecosystem functioning certain species may be considered redundant, while in view of evolutionary processes certain ecosystem processes may be considered redundant. Where do the two approaches meet and when should they be separated? This paper touches upon this question by referring to the dual hierarchy of ecological systems.