Forest Understorey Vegetation: Colonization and the Availability and Heterogeneity of Resources

Understorey vegetation comprises a major portion of plant diversity and contributes greatly to nutrient cycling and energy flow. This review examines the mechanisms involved in the response of understorey vegetation to stand development and the overstorey canopy following disturbances. The overall abundance and diversity of the understorey is enhanced with the availability and heterogeneity of light, soil nutrients, soil moisture, and substrates. Vascular plants are positively impacted by the availability and heterogeneity of light and soil nutrients, whereas non-vascular vegetation is more strongly influenced by colonization time, soil moisture, and substrates, and is decreased with a higher proportion of broadleaf overstorey. The availability of resources is a prominent driver toward the abundance and diversity of understorey vegetation, from the stand initiation to stem exclusion stage under a single-species dominated overstorey. However, resource heterogeneity dominates at the later stages of succession under a mixed overstorey. Climate and site conditions modify resource availability and heterogeneity in the understorey layer, but the extent of their influences requires more investigation. Forest management practices (clearcutting and partial harvesting) tend to increase light availability and heterogeneity, which facilitates the abundance and diversity of understorey vascular plants; however, these factors reduce the occurrence of non-vascular plants. Nevertheless, in the landscape context, anthropogenic disturbances homogenize environmental conditions and reduce beta-diversity, as well, the long-term effects of anthropogenic disturbances on understorey vegetation remain unclear, particularly compared with those in primary forests.