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Exclusion of wild ungulates is not the Holy Grail

KOvacs Bence erdos foto

A recent study authored with the significant contribution of the HUN-REN Centre for Ecological Research, Institute of Ecology and Botany was published with this title in the D1-ranked journal Journal of Environmental Management. Based on a field experiment, the authors demonstrate that under moderate ungulate density, the effect of forest management interventions on woody regeneration is stronger than the effect of ungulate exclusion per se. Over time, within fenced areas, the regeneration of the light-demanding sessile oak is hindered by faster-growing, more shade-tolerant competitors such as hornbeam, whereas outside the fences, under moderate browsing pressure, interspecific competition may be less intense. The seven-year study was initiated by Bence Tóth, Biology–Chemistry Teacher (MA) at ELTE Eötvös Loránd University, and Bence Kovács (Forest Ecology Research Group, Institute of Ecology and Botany, HUN-REN Centre for Ecological Research), and was later continued by Eszter Lilla Szabó, PhD student (Doctoral School of Biology, ELTE Eötvös Loránd University), under the supervision of Péter Ódor. The study highlights the importance of harmonizing forestry and game management practices to ensure the natural regenerative capacity of forests.

Managing natural regeneration in oak–hornbeam forests requires balancing light availability, competition dynamics, and herbivory pressure. Although fencing is often considered indispensable for protecting oak regeneration, our seven-year experiment shows that the influence of browsing exclusion is frequently overestimated. Silvicultural treatments, particularly those altering canopy openness, exerted a stronger effect on the early/mid-term sapling growth than fencing. Yet the study also highlights that maintaining low to moderate ungulate densities is crucial – not only for long-term regeneration success but even for sustaining the conditions under which silvicultural treatments can outperform browsing.

The researchers used the research infrastructure of the Pilis Forestry Systems Experiment to test the interactive effects of forestry and exclusion treatments. Within the framework of this experiment established in 2014, treatments of conventional rotation forestry (namely, micro clear-cutting, retention tree groups and preparation cutting) and of continuous cover forestry (gap-cutting) was compared to control plots in the closed, two-layered oak–hornbeam stand in the Pilis Mts. (Hosszú hill). For seven consecutive years, the researchers followed the browsing probability, survival, and growth of pairs of fenced and unfenced saplings selected and consolidated in the autumn of 2014.

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Silvicultural treatments drive early regeneration growth more strongly than browsing – but only when ungulate densities remain controlled

The applied (micro)clear-cutting (which essentially represent areas of final cutting) and artificial gaps resulted in the greatest increases in shoot growth and estimated leaf area across all tree species. This effect is primarily attributable to enhanced light availability and soil moisture conditions. In these treatments, the extent of browsing and overall ungulate impact temporarily increased – as expected given the increased food availability –; however, the survival of the selected individuals remained stable, because ungulate density in the studied landscape is moderate, largely due to the comprehensive game management practices of Pilisi Parkerdő Ltd. Under low to moderate browsing pressure, individuals outside fenced areas have a higher chance of escaping browsing pressure.

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According to Bence Kovács, the corresponding author of the article, it is important to emphasize that “this balance is fragile. The finding that forestry treatments had a stronger effect on individual growth depends on a controlled ungulate population.” In regions with higher densities and browsing levels comparable to those observed here (~60% of individuals browsed) can translate into severe growth reductions or regeneration failure. “Thus, the stronger effect of the treatments observed in this study does not mean that oak regeneration is better without fencing under the current landscape structure and browsing pressure, but rather demonstrates that browsing pressure can be managed if ungulate density remains within an ecologically compatible limits”, the researcher adds.

In the case of shrubs, however, forest management treatments and ungulate exclusion had effects of similar magnitude, which on the one hand indicates their sensitivity to browsing (a substantial difference in height growth between fenced and unfenced plots) and, on the other hand, highlights their role in shaping regeneration dynamics among tree species, especially of the target tree species.

Species-specific trajectories reveal competition as a dominant force

Oaks and manna ash benefited from initial browsing protection, but these advantages diminished due to intensified competition from faster-growing and typically shade-tolerant tree species – especially from hornbeam.

Hornbeam’s rapid juvenile growth and architectural plasticity allowed it to overtop oaks in both gaps and clear-cuts, and competitive effects were amplified inside exclosures. Consequently, in fenced areas where oak regeneration or restoration is a key objective, the suppression of competitor species is necessary, an area in which Pilisi Parkerdő Ltd. already has substantial practical experience. Based on the findings of the study, controlling competition is just as important as managing herbivory, and that fencing without subsequent tending is insufficient for ensuring long-term oak success under the framework of continuous cover forestry.

Shrubs, being highly palatable (and therefore most likely to be browsed) responded most strongly to ungulate exclusion, showing the largest differences in height growth between fenced and unfenced plots. As a result, their presence can substantially reduce browsing pressure on target tree species outside fenced areas and, and – under controlled ungulate densities – can indirectly facilitate oak regeneration.

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Gap-cutting can rival clear-cutting for oak regeneration

Medium-sized gaps (cca. 1:1 gap diameter to tree height ratio) generated microclimatic conditions that supported oak survival and growth comparable to clear-cuts. This result challenges assumptions that large openings or extensive fencing are necessary for oak regeneration and supports the feasibility of continuous cover forestry approaches. However, the success of gap-based regeneration still assumes ungulate densities low enough to prevent disproportionate browsing pressure.

Temporal shifts: from browsing limitation to competition limitation

A clear transition in growth-reducing factors emerged over the seven years:

  • Years 1–4: browsing and light availability shape height increment.
  • Years 5–: interspecific competition – especially from hornbeam – becomes the dominant constraint, particularly in fenced plots.

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This reinforces that ungulate exclusion alone is insufficient; regeneration ultimately hinges on managing competition for saplings to escape herbivore reach.

Management implications

The findings support an integrated, adaptive regeneration strategy:

  • Ungulate population control is fundamental. Only at low to moderate densities can browsing remain compatible with successful oak regeneration and allow silvicultural treatments to dominate growth responses.
  • Use moderate gap sizes to provide adequate light without excessively attracting herbivores.
  • Employ fencing strategically and temporarily – primarily to support the earliest regeneration stages.
  • Conduct targeted tending to reduce overtopping by shade-tolerant competitors.
  • Retain or support a shrub layer in the forest matrix, which contributes to browsing dilution and enhances stand structural complexity.

Our study demonstrates that while ungulate exclusion is not the Holy Grail, neither can natural regeneration succeed without effective ungulate population management. Silvicultural treatments exerted stronger early influence on growth, but this advantage was only possible because browsing pressure remained at ecologically compatible levels. Integrating population control, canopy management, and competition regulation provides a robust pathway for achieving resilient oak regeneration under continuous cover forestry across Central Europe.” concludes Bence Kovács.

Photo credits: Bence Tóth