Abstract
We present a conceptual model in which plant-soil interactions in grasslands are characterized by the extent to which water is limiting. Plant-soil interactions in dry grasslands, those dominated by water limitation ('belowground-dominance'), are fundamentally different from plant-soil interactions in subhumid grasslands, where resource limitations vary in time and space among water, nitrogen, and light ('indeterminate dominance'). In the belowground-dominance grasslands, the strong limitation of soil water leads to complete (though uneven) occupation of the soil by roots, but insufficient resources to support continuous aboveground plant cover. Discontinuous aboveground plant cover leads to strong biological and physical forces that result in the accumulation of soil materials beneath individual plants in resource islands. The degree of accumulation in these resource islands is strongly influenced by plant functional type (lifespan, growth form, root:shoot ratio, photosynthetic pathway), with the largest resource islands accumulating under perennial bunchgrasses. Resource islands develop over decadal time scales, but may be reduced to the level of bare ground following death of an individual plant in as little as 3 years. These resource islands may have a great deal of significance as an index of recovery from disturbance, an indicator of ecosystem stability or harbinger of desertification, or may be significant because of possible feedbacks to plant establishment. In the grasslands in which the dominant resource limiting plant community dynamics is indeterminate, plant cover is relatively continuous, and thus the major force in plant-soil interactions is related to the feedbacks among plant biomass production, litter quality and nutrient availability. With increasing precipitation, the over-riding importance of water as a limiting factor diminishes, and four other factors become important in determining plant community and ecosystem dynamics: soil nitrogen, herbivory, fire, and light. Thus, several different strategies for competing for resources are present in this portion of the gradient. These strategies are represented by different plant traits, for example root:shoot allocation, height and photosynthetic pathway type (C3 vs. C4) and nitrogen fixation, each of which has a different influence on litter quality and thus nutrient availability. Recent work has indicated that there are strong feedbacks between plant community structure, diversity, and soil attributes including nitrogen availability and carbon storage. Across both types of grasslands, there is strong evidence that human forces that alter plant community structure, such as invasions by nonnative annual plants or changes in grazing or fire regime, alters the pattern, quantity, and quality of soil organic matter in grassland ecosystems. The reverse influence of soils on plant communities is also strong; in turn, alterations of soil nutrient supply in grasslands can have major influences on plant species composition, plant diversity, and primary productivity.