Xylem traits, leaf longevity and growth phenology predict growth and mortality response to defoliation in northern...

Defoliation outbreaks are biological disturbances that alter tree growth and mortality in temperate forests. Trees respond to defoliation in many ways; some recover rapidly, while others decline gradually or die. Functional traits such as xylem anatomy, growth phenology or non-structural carbohydrate (NSC) storage could explain these responses, but idiosyncratic measures used by defoliation studies have frustrated efforts to generalize among species. Here, I test for functional differences with published growth and mortality data from 37 studies, including 24 tree species and 11 defoliators from North America and Eurasia. I synthesized data into standardized variables suitable for numerical models and used linear mixed-effects models to test the hypotheses that responses to defoliation vary among species and functional groups. Standardized data show that defoliation responses vary in shape and degree. Growth decreased linearly or curvilinearly, least in ring-porous Quercus and deciduous conifers (by 10-40% per 100% defoliation), whereas growth of diffuse-porous hardwoods and evergreen conifers declined by 40-100%. Mortality increased exponentially with defoliation, most rapidly for evergreen conifers, then diffuse-porous, then ring-porous species and deciduous conifers (Larix). Goodness-of-fit for functional-group models was strong (R2c = 0.61-0.88), if lower than species-specific mixed-models (R2c = 0.77-0.93), providing useful alternatives when species data are lacking. These responses are consistent with functional differences in leaf longevity, wood growth phenology and NSC storage. When defoliator activity lags behind wood-growth, either because xylem-growth precedes budburst (Quercus) or defoliator activity peaks later (sawflies on Larix), impacts on annual wood-growth will always be lower. Wood-growth phenology of diffuse-porous species and evergreen conifers coincides with defoliation and responds more drastically, and lower axial NSC storage makes them more vulnerable to mortality as stress accumulates. These functional differences in response apply in general to disturbances that cause spring defoliation and provide a framework that should be incorporated into forest growth and vegetation models.