Effects of twenty years of ungulate browsing on forest regeneration at Paneveggio reserve, Italy

Forest ecosystems are threatened by different natural disturbances. Among them, the irruption of large herbivores represents one of the most alarming issues. Several local-scale studies have been carried out to clarify the mechanisms governing ungulate-forest interactions, to understand the effect of wild ungulates overabundance, and to apply conservation plans. However, information at large scales, over long periods of observation and from unmanipulated conditions is still scarce. This study aims to improve our knowledge in this field by using repeated inventories to investigate: the types of damage produced by ungulate populations on young trees, the drivers that stimulate browsing activity and its consequences on the specific composition of seedlings and saplings. To reach these goals, we used data collected during a twenty-year monitoring program (1994-2014) in the forests of Paneveggio-Pale di San Martino Nature Park (Italy). We applied descriptive statistics to summarize the data, GLMs to identify the drivers of browsing activity and Non-Metric Multidimensional Scaling (nMDS) ordinations to investigate the changes in specific composition of young trees across 20 years. We detected increasing browsing activity from 1994 to 2008 and a decline in 2014. Ungulates browsed preferentially in mature stands, and fed mostly on seedlings and saplings under 150 cm of height. The analysis of the environmental drivers of browsing pressure on the smallest size classes of plants suggests that foraging behavior is influenced by snowpack conditions, ungulate density and seasonality. Moreover, results underline the fact that ungulates feed mostly on palatable species, especially European rowan, but can also use unpalatable plants as emergency food under high competition levels. nMDS results suggest that rowan seed dispersion might be promoted by deer movements, however, saplings of this species were not able to exceed 30 cm of height because of heavy browsing. This bottleneck effect led to the dominance of unpalatable species, mostly Norway spruce, reducing diversity during forest regeneration. If prolonged, this effect could lead to a reduction of tree species richness, with cascading effects on many parts of the ecosystem, and threatening the resilience of the forest to future disturbances.