Soil erosion under forest hampers beech growth: Impacts of understory vegetation degradation by sika deer

The overpopulation of large herbivorous mammals degrades understory vegetation and hence promotes soil erosion in some northern hemisphere forests. Topsoil removal via erosion reduces soil water and nitrogen (N) content as well as exposes tree roots. However, whether topsoil removal inhibits tree growth through water and N uptake limitations has not been examined. This study aimed to measure the vertical length from exposed roots surface to the soil surface (exposed root height; ERH) under degraded forest understory vegetation and to evaluate its influence on tree growth. Specifically, we examined the relationships between ERH and beech growth and identified the timing of growth reduction based on annual tree-ring analysis at the individual scale. Furthermore, we discussed the effects of water and N limitations on beech growth reduction. We studied 12 Japanese beech trees on Mt. Sanpo, southern Kyushu Island, Japan, where understory vegetation was degraded and lost due to herbivory by Japanese sika deer from 1980 to 2003. Lower amounts of leaf production and relative growth rates of stem basal area increment (BAI) were observed in beech trees with higher ERH. Segmented regression analysis, which detects the changing time of a temporal trend (i.e., break-point), revealed that the BAI has resulted in a decreasing trend after 1997.6 +/- 9.0 (yr), which coincided with the field obser-vations on understory degradation in the study site. After the break-point of the BAI, the BAI of each study tree was negatively related to intrinsic water use efficiency, an integrated index of water limitation. These results suggest that soil erosion after the degradation of understory vegetation reduced beech growth, possibly because of the water limitations associated with root exposure. Reduced beech growth will reduce the supply of the litter layer that prevents topsoil erosion, which may enhance further soil erosion and tree growth reduction.