Temporal patterns of ungulate herbivory and phenology of aspen regeneration and defense.

Ungulate herbivory can create strong top-down effects on forest recruitment, especially after fire. Defense strategies of tree species against ungulate herbivory include escape through vertical growth and resistance through the production of defense compounds. Using a four-way fence design and camera traps we characterized the differential impacts of ungulate herbivores (deer, elk, cattle) on aspen forest recruitment and plant defense responses and how they vary depending on the timing of herbivory. Aspen height growth was greatest between June and August and ungulate use of aspen was highest in July and August. Three years after fire, height of aspen differed among fence treatments with full ungulate exclusion > deer-only plots > native ungulate plots > fenceless plots: 108 ± 4 cm, 94 ± 4 cm, 89 ± 4 cm, and 65 ± 4 cm, respectively. Fenceless plots had the highest rates of removal of apical meristems by the end of 2014 and 2015 (61% and 53%, respectively). Native ungulate plots, and deer-only plots both had similar removal of apical meristems in 2014 (37% and 39%, respectively). The highest phenolic glycoside concentrations were associated with an 80% reduction in meristem removal and four-fold greater aspen height by the end of summer. Low nitrogen was associated with an 86% reduction in apical meristem removal and threefold greater aspen height. In conclusion, our study suggests that high ungulate abundance can have detrimental impacts on forest recruitment and that high aspen defense chemistry and lower leaf N deters ungulate herbivory, especially in the late summer.