Climatic effects on soil trophic networks and the resulting humus profiles in holm oak (Quercus rotundifolia) forests in the High Atlas of Morocco as revealed by correspondence analysis

Multivariate methods have been widely used for revealing the structures of communities, and in this paper we explore one particular method, namely correspondence analysis (also called reciprocal averaging), for studying humus profiles by the 'method of small volumes'. The present study was done on humus profiles under holm oak (Quercus rotundifolia), an evergreen Mediterranean species, in the High Atlas of Morocco. Three sites (1500 m, 1700 m, 1900 m altitude) and 2 years (1999 and 2002) were compared. The humus form is Dysmull (mull with thick litter horizons), with variations in the thickness of the OL (entire leaves), OF (fragmented leaves with faecal pellets) and A (hemorganic) horizons according to altitude and year. The dead leaves are rapidly incorporated into holorganic (earthworm, insect) and hemorganic (enchytraeid) animal faeces, which form the bulk of the OF and A horizons. The S horizon (weathering parent rock) shows the greatest development of the root system. As altitude increases more fresh litter (OL) or more humified organic matter (OF, A) is accumulated. Variation from year to year is depicted by opposite differences in the amount of entire oak leaves and of dead roots. Humus components (classes) are used as active (main) variables, after standardization of their means and variances. The addition of numerous passive (additional) variables, standardized in the same way as active variables, enabled us to understand the influence of biological and climatic effects on the composition of humus profiles and soil trophic networks.