The intensity of competition is a physiological concept, related directly to the well-being of individual organisms but only indirectly and conditionally to their fitness, and even more indirectly to the evolution of populations and the structure of communities. The importance of competition is primarily an ecological and evolutionary concept, related directly to the ecology and fitness of individuals but only indirectly to their physiological states. The intensity of competition is not necessarily correlated with the intensities of predation, disturbance, abiotic stress, or other ecological processes. The importance of competition is necessarily relative to the importances of other processes. Intensity refers primarily to the process of present competition, whereas importance refers primarily to the products of past competition. The distinction between the intensity and the importance of competition clarifies two long-standing ecological debates. Some ecologists have proposed that competition is greater in more stressful habitats, others the opposite, and still others that no such relationship exists. Evidence cited to refute or support these positions often confuses intensity and importance. Distinguishing between them focuses questions more sharply and indicates what sorts of new evidence should be sought. The more widely known debate over the prevalence of competition as an agent of community structure is a debate about the importance of competition, but evidence about the intensity of competition has often been used by both sides. We argue that intensity and importance need not be correlated, and so measurements of the intensity of competition are not directly relevant to this debate. This distinction also generates testable hypotheses and suggests directions for research. For example, we hypothesize that competition can be unimportant even if it is very intense: no such hypothesis is possible unless importance is distinguished from intensity. We discuss the application of these ideas to methods and theories used to study competition, ecological communities, and the evolution of competitive ability. We advocate a research approach that presumes multiple, interacting causes, including competition, affecting community structure, and we show how the distinction between intensity and importance helps to make this feasible.
We tested several hypotheses about the relationship of competition to abiotic stress, using the vegetation of the semiarid Piceance Basin of northwestern Colorado. We studied competition among the shrubs Amelanchier utahensis, Artemisia tridentata, and Symphoricarpos oreophilus, and between the trees Pinus edulis and Juniperus osteosperma, in 10 sites. We calculated several indices of biotic moisture stress, based on the slope, aspect, and evaluation of each site. Competition was measured by regression of the distance separating neighboring plants vs. the sum of their canopy areas. The slope of such a regression (if significant and positive) measures the intensity of competition, and its coefficient of determination (r2) measures the importance of competition, between the plants involved. We detected competition among these plants in all but one combination of species and in most sites. No significant differences in the intensity of competition were found within species combinations. Significant differences in the importance of competition were found in one of three interspecific combinations of shrub species, and in two of three combinations of tree species. Neither the intensity nor the importance of competition showed any consistent relationship with any index of abiotic moisture stress. Thus, no hypothesized relationship between abiotic stress and competition is supported. Our data also show no consistent relationship between the importance of competition and its intensity, supporting our hypothesis that the intensity and the importance of competition are independent.
The association between vegetation and soils from a geographically broad sampling of wetlands and adjoining uplands is reported for 38 hydric and 26 nonhydric soils, as recognized in the hydric soils list of the Soil Cortservation Service. Wetlands represented in the study include estuaries, pitcher plant bogs, prairie depressional wetlands, and western riparian lands. The agreement between vegetation and soils is clear with few exceptions. In general, hydric soils support hydrophytic plant communities, and nonhydfic soils support upland communities. Only 10% of the hydric soils sampled support upland communities and only 15 % of the nordlydric soils support wetland communities. Exceptions to the correspondence between vegetation and soils are discussed; local hydrology, the transitional nature of some soils, and other determinants of wetland vegetation structure (e.g., salinity, disturbance) seem to account for many of the observed discrepancies. A method that simplifies the complexity of soils and vegetation cannot be expected to represent accurately all details of their interrelations.
High elevation alpine and subalpine Rocky Mountain lakes in Colorado and southeastern Wyoming were examined to determine regional variability in water chemistry and their sensitivity to atmospheric deposition. Acid neutralizing capacity, pH, conductivity and concentrations of major anions and cations were compared. Regional differences in water chemistry are evident. The south-eastern most lakes have significantly higher pH, conductivity, ANC, and sums of acid and base concentrations than lakes in the other regions of the state. In contrast the northwestern most lakes are significantly more dilute than those from other regions. Despite these two regional differences, most regions are similar in having a wide range of variability in potential sensitivity of their lakes to acidification and nitrogen export. Many wilderness areas in western and eastern regions contains lakes that are extremely sensitive and other lakes not susceptible to deposition. Overall, 70% of the Colorado lakes are sensitive to acidification and 15% are extremely sensitive to acidification. All of the regions had lakes that are classified as susceptible or sensitive to acidification, with 12 of the 17 areas having all of their sampled lakes susceptible or sensitive. Generally NO 3 1À concentration in surface waters decreased from mid-season to late season; yet a large number of the lakes export NO 3 1À late in the season, suggesting nitrogen saturation. The results confirm the sensitivity of high elevation wilderness aquatic ecosystems in all regions of Colorado to acidification and nitrogen deposition.
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