Analyzing long-term impacts of ungulate herbivory on forest-recruitment dynamics at community and species level contrasting tree densities versus maximum heights

Abstract

Herbivores are constitutive elements of most terrestrial ecosystems. Understanding effects of herbivory on ecosystem dynamics is thus a major, albeit challenging task in community ecology. Effects of mammals on plant communities are typically explored by comparing plant densities or diversity in exclosure experiments. This might over-estimate long-term herbivore effects at community levels as early life stage mortality is driven by a multitude of factors. Addressing these challenges, we established a set of 100 pairs of ungulate exclosures and unfenced control plots (25 m2) in mixed montane forests in the Alps in 1989 covering a forest area of 90 km2. Investigations ran until 2013. Analogous to the gap-maker–gap-filler approach, dynamically recording the height of the largest trees per tree species in paired plots with and without exclosures might allow for assessing herbivore impacts on those individuals with a high probability of attaining reproductive stages. We thus tested if recording maximum heights of regenerating trees would better reflect effects of ungulate herbivory on long-term dynamics of tree regeneration than recording of stem density, and if species dominance patterns would shift over time. For quantifying the effects of ungulate herbivory simultaneously at community and species level we used principle response curves (PRC). PRCs yielded traceable results both at community and species level. Trajectories of maximum heights yielded significant results contrary to trajectories of total stem density. Response patterns of tree species were not uniform over time: e.g., both Norway spruce and European larch switched in their response to fencing. Fencing explained about 3% of the variance of maximum tree heights after nine years but increased to about 10% after 24 years thus confirming the importance of long-term surveys. Maximum height dynamics of tree species, addressed in our study, can thus reflect local dominance of tree species via asymmetric plant competition. Such effects, both within and among forest patches, can accrue over time shaping forest structure and composition.