Influence of Tree Species and Size on Bark Browsing by Large Wild Herbivores.

Although an important part of the ecosystem, large wild herbivores (LWH), especially red deer (Cervus elaphus L.), cause significant damage to economically valuable timber in forests of Central Europe. Recent work has demonstrated that less valuable softwood broadleaved trees can act as "biological control" that helps reduce bark browsing on more valuable trees in a mixed stand. To better understand the factors that influence how much bark area and mass are removed by LWH from these broadleaved trees, we took advantage of a novel "natural" experiment that occurred after a breach in a herbivory exclosure surrounding a 10-year old mixed broadleaved/conifer stand in the Western Carpathians in north-western Slovakia. We measured the area of old (up to 2 years previously) and new browsed patches on stems of common aspen (Populus tremula L.), common rowan (Sorbus aucuparia L.) and goat willow (Salix caprea L.), and their position along the vertical profile of the stem. The browsed bark area (cm2) was then converted to the bark mass (g) removed and the proportion of browsed bark to total bark (%) using conversion equations. Our models demonstrated that the amount of bark removed was influenced by tree species, stem diameter, age of browsing (old vs. new), and stem section along the vertical profile. LWH removed the most bark area from willow but the most bark mass from aspen because aspen had thicker bark than the other tree species. Bark browsing was greater on trees > 6 cm basal diameter. The distribution of bark browsing along the vertical profile was symmetrical (unimodal) with maximum intensity at 101−125 cm from the ground, which corresponds with the height most optimal for feeding by red deer. However, previous browsing in 2019 and 2020 caused new browsing on willow in 2021 to be focused in stem sections lower (51−75 cm) and higher (126−150 cm) than that optima. By quantifying browsing patterns and the amount of bark that is accessible to LWH for forage on the most attractive softwood broadleaved trees, our work will contribute to developing better methods for protecting commercially important species such as European beech (Fagus sylvatica L.) and Norway spruce (Picea abies L. Karst.) in areas of Central Europe that are greatly affected by increasing population density of LWH, especially red deer.