Interacting effects of insect and ungulate herbivory on Scots pine growth

Most plants are subjected to damage from multiple species of herbivores, and the combined impact on plant growth can be non-additive. Since plant response to herbivores tends to be species specific, and change with repeated damage, the outcome likely depend on the sequence and number of attacks. There is a high likelihood of non-additive effects on plant growth by damage from mammals and insects, as mammalian herbivory can alter insect herbivore damage levels, yet few studies have explored this. We report the growth response of young Scots pine trees to sequential mammal and insect herbivory, varying the sequence and number of damage events, using an ungulate-pine-sawfly system. Combined sawfly and ungulate herbivory had both additive and non-additive effects on pine growth—the growth response depended on the combination of ungulate browsing and sawfly defoliation (significant interaction effect). Repeated sawfly herbivory reduced growth (compared to single defoliation) on un-browsed trees. However, on browsed trees, depending on when sawfly defoliation was combined with browsing, trees exposed to repeated sawfly herbivory had both higher, lower and the same growth as trees exposed to a single defoliation event. We conclude that the sequence of attacks by multiple herbivores determine plant growth response.

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

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.

Herbivory modulates soil CO2 fluxes after windthrow: a case study in temperate mountain forests

Ungulate herbivory can alter functional plant communities of early-successional forest ecosystems. The consequences of such vegetation changes on soil carbon cycling are still not fully understood. Here, we used an ungulate exclusion experiment to investigate how different levels of herbivory and associated changes in vegetation succession modulate soil CO2 efflux and its heterotrophic and autotrophic sources following windthrow in temperate mountain forests. Our results indicate that only high levels of ungulate herbivory and associated vegetation shifts from tree to rather grass dominated plant communities affect soil CO2 fluxes. We did not find evidence that a moderate herbivory level and accompanied smaller shifts in the functional plant community affect soil CO2 fluxes. A greater soil CO2 efflux under the influence of high herbivory pressure was primarily attributed to accelerated heterotrophic respiration, likely due to warmer soil conditions. Moreover, autotrophic respiration from grass roots and associated microbial communities is suggested to contribute to higher soil CO2 fluxes. We conclude that intense herbivory and accompanied successional changes in the functional plant community enhance soil carbon losses following forest windthrow. This might have negative consequences for the soil carbon stocks and for the climate system.