Valleys and Hillslopes: A Geomorphic Foundation for Landscape Ecology

Moisture-nutrient gradients have been found to be the most important environmental gradients determining the distribution and composition of plant communities. Landscapes on which plant communities exist are composed of valleys and ridgelines, with hillslopes in between them. Since water flow paths are directed down slopes, processes determining hillslope morphology and arrangement play an essential role in plant community organization and dynamics. Hillslope morphology, substrate characteristics and climate determine flow routing and water budgets along slopes. Wetness is a function of transmissivity, contributing area and slope gradient. Movement of nutrients along hillslopes generally follows wetness values, and is affected by soil type. Plant species have different tolerances to wetness and nutrients; hillslope length and slope angle determine the moisture-nutrient gradient, and in turn the shape of plant tolerance curves. Temporal scales required for significant topographic change along hillslopes may often be long compared to those for plant community dynamics. When considered in landscape ecology, hillslope shape and arrangement are thus often considered constants. Although landscape morphology may change over time and among different regions (with tectonic, geomorphic and climatic processes leaving their imprints on landscapes), an attempt has been made in the literature to put forth robust topographic scaling relations. This paper, using a series of examples, explores connections between landscape structure and plant communities. For example, Hack's law states that drainage basins become more elongate as area increases. This implies that basins should have approximately the same proportion of landscape in each hillslope position, suggesting some constancy in contributing area patterns for hillslopes in different-sized basins. Distributions of wetness values and plant population tolerance curves seem to confirm this for smaller basins. Hillslope length and steepness are related to drainage density and relative relief. Various studies have sought relations between drainage density and slope gradient; the latter is a determinant of wetness values. Studies have found both negative and positive correlations between drainage density and slope gradient. The nature of hillslope processes (e.g., overland flow vs. mass wasting dominated, or quickly eroding vs. slowly eroding landscapes) has been used to explain the correlation. It has also been suggested that the degree of channelization may be important in determining slope steepness. Plant species respond to steeper slopes by having narrower tolerance curves and less overlap with other species. This has important implications for biodiversity and plant community organization.