The flora of the deciduous forests at the base of the north Estonian limestone escarpment is species rich, with an exceptionally high number of rare bryophyte species. Relationships between species richness of bryophyte and herb layers and biotic and environmental conditions were studied, using General Linear Mixed Models. Human disturbance (waste deposit, tree damage etc) was significantly negatively correlated with species richness of both plant layers. Soil nitrogen content was negatively and soil water retention positively correlated with bryophyte species richness, while herb richness was unrelated to soil factors. After eliminating the effects of environment, negative correlations in species richness and cover between the bryophyte and herb layers were discovered on finer scales (1 m²), referring to biotic interactions. This relationship was obscured with the simple correlation analysis. On the other hand, the positive correlation in species pools between the bryophyte and herb layers (0.1 ha) was insignificant. The species pools of both bryophyte and herb layers were significantly positively correlated with the species richness of the tree layer. In summary, bryophyte and herb layer richness responded differently to environmental conditions, but human disturbance significantly decreased the richness of both layers. Due to the uniqueness and small area of these forests we recommend protection and restoration of disturbed sites.

Nomenclature: Ingerpuu et al. (1994) for bryophytes, Leht (1999) for phanerogams.

Abbreviation: GLMM = General Linear Mixed Model.