Abstract:

Aims Many past practices in afforestation and forest management were instrumented for addressing the issues of tree species selection, planting regimes and development of overstory structure, but neglected understory vegetation. Our objective was to determine the controlling factors of herb-layer plant distribution and the importance of topography in determining local-scale spatial patterns of herbaceous plants.
Methods The occurrence and distribution of herb-layer plants were investigated on 26 plots in the Xiaoshegou catchment of Lingkong Mountain, Shanxi Province, China. Community types were classified using two-way indicator species analysis (TWINSPAN) and the relationship between the distribution and abundance of herb-layer species and environmental gradients was analyzed using the canonical correspondence analysis (CCA) ordination method. Forward selection and Monte Carlo permutation test were used to select the factors important in determining the herb-layer plant distribution. Partial CCA (PCCA) was also performed to partition the variance that was explainable by categorical habitat and biotic factors studied.
Important findings The 26 survey plots were classified into six groups characterized by the dominant overstory tree species. The results of CCA ordination reflected the relationship between herb-layer community structure and selective environmental variables. The classification of 26 plots in CCA ordination was consistent with the result of TWINSPAN. Forward selection and Monte Carlo test suggested that stand type, soil nutrients and slope position were the most important factors determining understory plant distribution. PCCA revealed that habitat and biotic factors together explained 42.9% of variance in the distribution of understory herbaceous plants. Habitat factors alone explained 31.8% of the variance, biotic factors alone explained 7.8% of the variance and interaction between habitat and biotic factors explained 3.2% of the variance. The partitioning of variance using the PCCA helped with identifying the important habitat factors regulating understory herbaceous plant distribution at the study site. However, the fact that more than half of the variance was unaccounted for by the factors studied suggests that other factors we did not measure could also play a role in determining the occurrence and distribution of herbaceous plants on the forest floor, e.g., human activities and random events. Our study demonstrates the importance of both topography and overstory tree species in determining the occurrence and distribution of herb-layer plant species in temperate forest of mountainous areas.

Key words: canonical correspondence analysis (CCA), temperate forest, understory communities, topography, variance partitioning, vegetation-site relationship