Showing papers by "James B. Grace published in 1999"

The factors controlling species density in herbaceous plant communities: an assessment

Abstract: This paper evaluates both the ideas and empirical evidence pertaining to the control of species density in herbaceous plant communities. While most theoretical discussions of species density have emphasized the importance of habitat productivity and disturbance regimes, many other factors (e.g. species pools, plant litter accumulation, plant morphology) have been proposed to be important. A review of literature presenting observations on the density of species in small plots (in the vicinity of a few square meters or less), as well as experimental studies, suggests several generalizations: (1) Available data are consistent with an underlying unimodal relationship between species density and total community biomass. While variance in species density is often poorly explained by predictor variables, there is strong evidence that high levels of community biomass are antagonistic to high species density. (2) Community biomass is just one of several factors affecting variations in species density. Multivariate analyses typically explain more than twice as much variance in species density as can be explained by community biomass alone. (3) Disturbance has important and sometimes complex effects on species density. In general, the evidence is consistent with the intermediate disturbance hypothesis but exceptions exist and effects can be complex. (4) Gradients in the species pool can have important influences on patterns of species density. Evidence is mounting that a considerable amount of the observed variability in species density within a landscape or region may result from environmental effects on the species pool. (5) Several additional factors deserve greater consideration, including time lags, species composition, plant morphology, plant density and soil microbial effects. Based on the available evidence, a conceptual model of the primary factors controlling species density is presented here. This model suggests that species density is controlled by the effects of disturbance, total community biomass, colonization, the species pool and spatial heterogeneity. The structure of the model leads to two main expectations: (1) while community biomass is important, multivariate approaches will be required to understand patterns of variation in species density, and (2) species density will be more highly correlated with light penetration to the soil surface, than with above-ground biomass, and even less well correlated with plant growth rates (productivity) or habitat fertility. At present, data are insufficient to evaluate the relative importance of the processes controlling species density. Much more work is needed if we are to adequately predict the effects of environmental changes on plant communities and species diversity.

The relationship between species density and community biomass in grazed and ungrazed coastal meadows

Abstract: Previous studies have indicated that the relationship between community biomass and species density can be represented by a multivariate model in which abiotic variables influence species density both through effects on biomass and through effects on the species pool. In this paper, we use data from grazed and ungrazed coastal meadows in Finland to evaluate and extend this general conceptual model of the factors controlling species density. Structural equation analysis was used to evaluate a model for all meadows and then to perform a multigroup analysis to determine how grazed and ungrazed meadows differ. By itself, biomass could explain only 12% of the variation in species density while the multivariate model was able to explain 47% using five types of predictor variables: site, soil, flooding, grazing, and biomass. Analyses found that flooding explained the greatest amount of variability in species density, primarily through negative effects on the species pool. Grazing was also found to have a strong effect on species density and results suggest that its negative influence may be largely through reductions in the species pool in grazed meadows. The most important difference found between grazed and ungrazed meadows was that species density had a strong negative relationship to biomass in the ungrazed meadows but no significant relationship in the grazed ones. Thus, it appears that the influence of competition on species density was much greater in ungrazed meadows compared to grazed ones.

Effects of environmental change on plant species density: comparing predictions with experiments

Abstract: Ideally, general ecological relationships may be used to predict responses of natural communities to environmental change, but few attempts have been made to determine the reliability of predictions based on descriptive data. Using a previously pub- lished structural equation model (SEM) of descriptive data from a coastal marsh landscape, we compared these predictions against observed changes in plant species density resulting from field experiments (manipulations of soil fertility, flooding, salinity, and mammalian herbivory) in two areas within the same marsh. In general, observed experimental responses were fairly consistent with predictions. The largest discrepancy occurred when sods were transplanted from high- to low-salinity sites and herbivores selectively consumed a particularly palatable plant species in the transplanted sods. Individual plot responses to some treatments were predicted more ac- curately than others. Individual fertilized plot responses were not consistent with predictions (P > 0.05), nor were fenced plots (herbivore exclosures; R2= 0.15) compared to unfenced plots (R2 = 0.53). For the remaining treatments, predictions reasonably matched responses (R2 = 0.63). We constructed an SEM for the experimental data; it explained 60% of the variance in species density and showed that fencing and fertilization led to decreases in species density that were not predicted from treatment effects on community biomass or observed distur- bance levels. These treatments may have affected the ratio of live to dead biomass, and competitive exclusion likely decreased species density in fenced and fertilized plots. We conclude that experimental validation is required to determine the predictive value of comparative relationships derived from descriptive data.

The effects of landscape position on plant species density: Evidence of past environmental effects in a coastal wetland

Abstract: Here we propose that an important cause of variation in species density may be prior environmental conditions that continue to influence current patterns. In this paper we investigated the degree t...