Density-related effects of deer browsing on the regeneration dynamics of boreal forests

1. The density of large herbivores is a major driver of forest ecosystem structure and function in conjunction with episodic disturbances, especially in forests with a regeneration strategy based on shade-tolerant seedlings capable of re-establishing canopy dominance (advance regeneration). Yet, uncertainty about the relationships between forest regeneration, herbivore density and other disturbances makes it difficult to set population goals. Using an innovative controlled browsing experiment, we investigated the relationships between the regeneration dynamics of balsam fir Abies balsamea, the density of white-tailed deer Odocoileus virginianus and timber harvesting. 2. We hypothesize that advance tree regeneration either: (i) recovers approximately linearly as deer density is reduced; (ii) recovers exponentially; or (iii) does not recover because factors other than browsing control advance regeneration. We tested these alternatives through manipulation of deer densities (0, 7.5, 15 deer km(-2) and in situ local densities) and forest cover (clearcut and uncut forest). 3. Balsam fir seedling mortality decreased exponentially with decreasing deer density in clearcut and approximately linearly in uncut forest. Independently of deer density, the recruitment of seedlings in clearcut dropped from 56 +/- 5% to 7 +/- 1% within 3 years. 4. Seedling growth increased exponentially with decreasing deer density in clearcut whereas no height growth was observed in uncut forest. 5. Overall, the abundance of fir saplings recovered exponentially in clearcut but remained low and independent of deer density in uncut forest. The abundance of spruce Picea spp. saplings was unrelated to deer density and increased with time. 6. Synthesis and applications. Forest disturbance from selective browsing at high deer densities over an extended period of time leads to recruitment failure following a canopy disturbance such as a clearcut. Indirect competitive advantage given to species resistant to browsing can shift forest composition. Nonlinear relationships between fir regeneration and deer densities imply that the level of culling required to reach herbivore densities compatible with natural regeneration of native forest is larger than expected if tree regeneration was proportional to deer density. In the boreal forest of Anticosti Island, local densities < 15 deer km(-2) achieved within 3 years following clearcut are compatible with the maintenance of native forest.