Ungulate herbivory alters leaf functional traits and recruitment of regenerating aspen

Herbivory by ungulates can affect forest regeneration success, but its long-term impacts on tree function and recruitment are less studied. We evaluated strategies of resistance, tolerance and vertical escape against ungulate herbivory by evaluating leaf traits (photosynthesis, morphology and chemistry) and growth rates of aspen in the presence and absence of ungulate herbivores 1, 2, 3 and 26 years after fires initiated aspen suckering. Over the initial 3-year period, ~60% of aspen stems in unfenced plots showed evidence of being browsed by ungulates. After 3 years, aspen in unfenced plots had smaller leaves, were 50% shorter, and had 33% lower nonstructural carbohydrate concentrations and 33% greater concentrations of condensed tannins, when compared with fenced aspen. Aspen exposed to ungulate herbivory over a 26-year period maintained smaller leaves, had lower annual radial growth rates and were still below the critical height threshold of 2 m required to escape ungulate herbivory for successful recruitment. In contrast, the average height of aspen protected from ungulates was approaching 6 m. Over the 26-year period leaves in unfenced plots had 41% lower nonstructural carbohydrate concentrations and greater expression of defense compounds-condensed tannins (63%) and phenolic glycosides (102%)-than leaves in fenced plots. Photosynthetic rates were slightly higher in aspen that experienced ungulate browsing, suggesting that changes in leaf anatomy and chemistry due to ungulate herbivory did not interfere with photosynthesis. Our data suggest that ungulate browsing increases investment in chemical defense, lowers nonstructural carbohydrate concentrations and reduces leaf area, which decreases the recruitment potential of regenerating aspen.