Interactions between earthworms, litter and trees in an old-growth beech forest

Forty plots were selected in an old-growth beech forest (Biological Reserve of La Tillaie, Fontainebleau State Forest, France), to embrace the whole range of site conditions and phases of vegetation dynamics. Soils are sandy, thus the nutrient status of the topsoil is very poor except when trees have access to an underlying limestone layer. The study was focused on the role of calcium in the sustainability of the beech ecosystem. Calcium is mostly redistributed through leaf litter accretion and the activity of litter-consuming organisms, but other sources are fallen wood and uprooted mounds. In each of the 40 plots, earthworm species were sampled, and measurements were taken in order to describe humus profiles, growth of adult trees, litter quantity and quality, and access to lime. Densities of soil-dwelling earthworms, calcium content of beech leaf litter, height of tallest trees and depth of the limestone layer were correlated, indicating a gradient of soil fertility which mainly results from long-term interactions between soil organisms and trees in varying geological conditions. Possible causal relationships and implications of calcium turnover for nature conservation were discussed in the light of existing knowledge.