The heterogeneity of humus profiles and earthworm communities in a virgin beech forest

Abstract

Thirty sites, encompassing a range of soil and vegetation conditions in the biological reserve of La Tillaie (Fontainebleau Forest, France) were investigated in April 1992. Beech (Fagus sylvatica L.) was the dominant tree species, with several developmental phases forming the forest patchwork. Sessile oak [Quercus petraea (Mattus.) Liebl.] was present but only as old relictual individuals. Gaps in the canopy cover were abundant, mostly caused by wind storms 2 years previously. The next most recent storm was 25 years before, resulting in distinct patches of full-grown trees. Humus profiles were classified and compared with the distribution of earthworm communities, canopy cover, and soil types. Geomorphology was responsible for the main part of the observed variation. Absence of lime in the substrate and direct contact with a sandstone stratum near the ground surface was associated with the absence of earthworms and the appearance of an OH horizon (moder humus). Elsewhere, earthworms were present and humus profiles did not display any OH horizon (mull or mull-like moder humus), but species composition was variable and strongly influenced by the thickness of the superficial sand deposit overlying limestone. On a thick (1 m or more) sandy substrate earthworm communities were dominated by epigeic species together with the anecic Lumbricus terrestris L. The species richness was higher on a shallower sandy substrate (50 cm) where lime was more accessible to tree roots and burrowing animals. The influence of the forest cycle of beech was visible in the latter case (covering most of the area), with an increase in the thickness of the OL and OF horizons and a decrease in endogeic earthworm populations during the phase of intense growth of beech. This fall in burrowing activity was apparent in gaps created by wind storms and fungal diseases within mature stands as early as 2 years after the fall of the trees.