Even low levels of spruce budworm defoliation affect mortality and ingrowth but net growth is more driven by competition

Defoliation reduces the growth and survival of trees, but this influence can be difficult to evaluate largely because of its interplay with various stand and site factors, especially for the highly dynamic defoliation of spruce-fir (Picea-Abies) forests caused by spruce budworm (SBW; Choristoneura fumiferana (Clem.)), the primary tree defoliator in North America. In this study, we developed statistical models to evaluate the influence of SBW defoliation, while considering its interaction with various stand and site factors, on spruce-fir stand dynamics of annual volume net growth, mortality, and ingrowth. The data were collected at intervals of 1 to 3 years from 560 permanent sample plots during the last SBW outbreak (1970s-1980s) in Maine, USA, and New Brunswick, Canada. These data comprise a wide range of observations of cumulative defoliation, especially at relatively low levels, that have been largely overlooked in previous studies. Our results strongly demonstrated that even relatively low levels of cumulative defoliation were significantly related to stand-level mortality and ingrowth, while net growth was more competition driven. Additionally, these stand dynamics were found to be not significantly affected by any of the site factors evaluated. These findings were consistent for Maine and New Brunswick despite their differences in forest management and SBW outbreak histories.