Protection of understory vegetation by deer exclosure fences prevent the reduction of beech growth due to soil erosion

Overbrowsing by ungulates has degraded understory vegetation in worldwide forests. Understory degradation causes soil erosion and environmental degradation, causing mature tree decline. Although ungulate exclosure fences are installed to protect vegetation, their effectiveness in preventing tree decline associated with soil degradation remains unclear. We aimed to fill this gap on slopes of Mt. Shiraga, Japan, where sika deer have overbrowsed for >20 years. Tree-ring records in 1960-2022 for eight beeches per each of inside and outside fences were analyzed to compare stem basal area increment (BAI, cm^-2 yr^-1) and trend change timings for BAI (i.e., break-point). Soil properties, exposed root height (ERH) as cumulative soil erosion index, and intrinsic water use efficiency (iWUE) as water limitation proxy were also compared. Compared with areas inside fences, areas outside fences had no understory vegetation, higher ERH, and higher soil hardness and moisture. BAI outside fences turned their growth trend from positive to negative since 2004, but not inside fences. Break-point of BAI outside fences matched the understory degradation period in the field. The long-term BAI trend (i.e., regression slope of BAI) after the break-point was negatively related to ERH but not to soil properties (e.g., soil hardness and moisture). Furthermore, BAI outside fences was negatively related to iWUE after the break-point, whereas BAI inside fences was not. These results indicate that the maintenance of understory by fencing prevents overbrowsing-associated mature tree decline through mitigating root exposure and related water limitation, despite the potential of water competition with understory.